ENVIS Forestry Bulletin

     Vol.8,No.1; YEAR-2008

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FOREST SCIENCE AND TECHNOLOGY SPECIAL
Seed Technology: Its Role in Conservation of Forest Genetic Resources
Seed Source Variation in Some Important Native Tree Species if Garhwal Himalaya
Environmental Ups and Downs
Significance of Seed Pathology in Forest Trees
Viewpoint: Implementation of Forest Tree Seed Certification and Legislation in India- What Are We Waiting For?
Quality Seed Production Through High Density Seed Orchards
Quality Seed Production of Grasses and Legumes for Forest Grazing Reserves and Derived Grazing Resources in India
Evaluation of Seed Quality During Maturation for Effective Collection
Pattern of Seed Dispersal and Regeneration in Dipterocarpus indicus: An Endemic Tree Species of Western Ghats
Seed Germination Studies in Different Seed Sources of Azadirachta indica for Higher Germination Percentage and Longer Viability
Seed Studies on Three Ficus Tree Species: Important Keystone Plant Resources
Desiccation and Temperature Tolerance of Recalcitrant Seeds- Madhuca Longifolia, Calophyllum and Antocarpus heterophyllus
Effect of Pretreatment and Storage on Germination Behaviour of Seeds of Plantago Ovata and Withania Somnifera
Variation in Germination and Seedling survival among uttrakhand Accessions of Jatropha curcas
Role of Seed Technology in Tree Improvement Programmes
Bibliography on Tree Seed Science and technology
Milestone: Robert Scott Troup
   
  ENVIS CENTRE ON FORESTRY
  FOREST RESEARCH INSTITUTE
  (INDIAN COUNCIL OF FORESTRY RESEARCH AND EDUCATION)
  DEHRADUN

SEED TECHNOLOGY: ITS ROLE IN CONSERVATION OF FOREST GENETIC RESOURCES

Manisha Tapliyal
Forest Research Institute, Dehradun-248006


Introduction

T ropical forests are the depleting wealth of diverse and valuable biodiversity with rare, threatened, endangered or otherwise valuable medicinal and commercially important tree species that need immediate conservation for posterity and as future scientific material for improvement and genetic engineering. Loss of biodiversity due to climate change or habitat destruction is gaining in political attention. An example of the increased attention for this topic is the establishment of a crop seed safety-storage facility at Svalbard (Spitsbergen), Norway (ABC Science, 2008). At present, gene banks store approximately 6.5 million seed samples (mostly crops). As a result of their limited budgets, most gene banks focus their resources mainly on efficient representation of the genetic diversity and description of the collection. Less attention tends to be paid to the physiological and health status of the seeds and to (temporary) seed storage conditions. A case study conducted at the Centre for Genetic Resources in the Netherlands showed that considerable improvements can be made in these areas, resulting in prolonged shelf life of the stored seed samples.

The report of a FAO/UNEP pilot study conducted about three decades back recommended the complementary role of ex situ seed banks in forest genetic resource conservation and identified the need to establish research programmes especially in the testing, storage and regeneration of tropical-forest tree seeds (Roche, 1975). This greatly increased interest in tree seeds, which caused high demands for seedlings of different tree species for planting during good and lean seed years. To meet this requirement, basic knowledge about seed biology and technology such as proper seed maturation time, harvesting, processing, germination, dormancy, viability and vigour and storage physiology for various tropical species should be known. Till date, much of the work has been done on temperate species and research on tropical and sub-tropical species has not matched those levels which has lot of scope for research in these areas.

In India, the National Genebank of National Bureau of Plant Genetic Resources (NBPGR) is the third richest in germplasm holding after China and USA and has largest infrastructure capacity to hold around 0.75 million base collections. But their seed bank collections include cereals, legumes, oilseeds, fiber crops, medicinal and aromatic plants, spices and condiments, millets and agroforestry species and there is very little representation of forestry species.

Establishment of the gene banks i.e. ex situ conservation, is one of the most effective biological techniques to conserve biodiversity. Conventional seed storage is believed to be a safe, effective and inexpensive method of ex situ conservation of plant genetic resources, which not only maintains its viability but its vigour too. An ideal condition to prolong the seed longevity is mainly dependent on seed moisture content, temperature and type of container used for storing the seed. To fulfill the conservation roles, seed-storage life must exceed the natural interval between germination and seed production for the next generation. Successful long term preservation is dependent on continuous viability monitoring with re-collection or regeneration, whenever viability drops below a minimum level.

Genetic erosion in a gene bank, i.e. loss of genetic material while under storage is one of the most concerning issues during conservation in recent years (Food and Agriculture Organisation of the United Nation, 1998). The cause is attributed mainly to the lack of operational resources. Research on conservation methodology and gene-bank management is, therefore, critical to streamline the operation of a genebank and to make it more cost-effective and attractive in funding. Little information is available on the loss and shift in genetic variability of an accession during each cycle of seed regeneration. The exact plant population required during seed regeneration in order to retain most of the original diversity is not known for most species. Also, a quick, low-cost, accurate seed vigour test for predicting seed quality for use in planning and programming of seed viability monitoring interval during storage is lacking.

Forest tree seeds remain the most popular and common source of reproductive propagules in large-scale afforestation and re-forestation programmes. Despite large-scale use, seed technologies for many species remain to be developed or are still inadequate. It is high time to explore the storage physiology of valuable tropical forest species and gather scientific knowledge for their cost-effective long term conservation through seed gene banks. Seed gene banks so far conserve only orthodox seeds, as genetic resource conservation diversifies into other species of tropical rain forests, where many of them have recalcitrant seeds, the problems and techniques would be different and would have to be developed for such species. In-vitro slow-growth and cryopreservation techniques are the most promising conservation methods applied to many recalcitrant seeds (Engelmann and Takagi, 2000).

Basic Gene Bank Design

The basic gene bank design consists of a seed storage facility, a seed drying system, a seed laboratory for seed processing and testing, a germplasm research laboratory, a packaging-cum-distribution laboratory and a data documentation laboratory. The principle in the design of a gene-bank storage system is the stepwise removal of outside air moisture and temperature, which is accomplished by the packaging room operating at 20 ± 2oC and 40 per cent R.H. as an anteroom for the slow dryer (15 ± 2oC and 15 per cent R.H.) and temporary seed cooler (15 ± 2oC and 40 per cent R.H.). The step-wise removal of air moisture allows the frost-free requirement of the long-term seed freezer to be achieved more easily.

Seed Science and Technology in Gene Banks

Knowledge and skills in seed science and technology are the foundation to run an efficient gene bank. From germplasm acquisition to seed multiplication and seed storage to germplasm distribution, seed science and technology knowledge and skills have to be applied when making decisions. However, forest genetic resources conservation is multidisciplinary and requires adequate knowledge of not only seed science but also forestry, plant genetics, physiology, protection, management, etc. for smooth functioning of seed gene banks.

The physical and physiological quality of seed lot accession has to meet the minimum standards of the genebank, otherwise seed regeneration has to be repeated (FAO/IPGRI. 1994). These standards include:

  • genetic quality – to ensure that the original genetic composition of an accession is maintained;

  • physiological quality – to ensure that the physiological state of a seed lot is optimum for maximum storage period;

  • physical quality – to ensure that a seed lot has no mechanical injuries thus affecting its storage life and also free from other extraneous matters; and

  • seed health quality – to ensure that a seed lot is not carrying infectious diseases, pests and weeds.

  • Genetic Quality
    Seed genetic quality in germplasm conservation means minimal changes in the genetic composition of an accession during seed regeneration and conservation. The changes could be due to the loss in variability during seed regeneration as a result of either genetic drift and conscious or unconscious selection.

    Physiological Quality
    The physiological quality of a seed lot determines its storage life and vigour. The two most important factors influencing this parameter are the state of seed physiological maturity during seed development and the degree of seed weathering incurred after seed physiological maturity. Seed weathering is the effect of injuries on seed due to extreme weather conditions such as too high moisture and high or low temperature extremes causing the affected seed to deteriorate at a quick rate. Response of seeds to storage conditions depends on the quality of seeds collected. Other than seed maturity, emptiness also affects the quality of seeds. Intermittent seeding patterns and high incidence of empty (aborted) seeds is widespread in tree species, the causes of which are yet to be fully understood.

    Physical Quality
    Physical seed quality in genetic resource conservation is mainly related to mechanical damage during seed harvesting, drying and processing. Sound and filled seeds must be separated from empty and non-viable seeds and inert fragments before storage. The seed must also be free of other alien material like soil, dust and small stones, noxious weeds and other damaged and broken seeds. This can be accomplished by making use of the physical properties like floating velocity, thickness, surface structure, colour, weight, length and width of the seed through the process of floatation, winnowing, seed blowing, aspiration, etc. Cleaning process has been made quick and effective by the modern cleaning machines.

    Health Quality
    Germplasm in a seed bank should be free of any pest . A plant germplasm may have assembly of tree genetic resources from different parts of the world. If the quarantine procedure is not meticulous during germplasm introduction, a collection of plant pathogens may also be assembled, which may intercross and establish during seed regeneration trial. Stringent plant quarantine regulations and protocols are in place throughout the world by national plant quarantine authorities and the International Plant Protection Convention of FAO (Food and Agriculture Organisation of the United Nation, 1997).

    Seed gene banks are mandated to conserve and distribute the widest representation (covering all possible variations), so its functions include germplasm acquisition, regeneration, conservation, evaluation, documentation and distribution. All these sequential procedures demand sound knowledge and skills of seed science and technology among gene-bank personnel. The following section briefly describes the close connection between plant germplasm conservation and seed science discipline which has also been demonstrated in Table 1.

    Table 1.Plant genetic resource conservation and distribution procedures in relation to seed science and technology

    Gene Bank Procedure Seed Science and Technology Knowledge
    Acquition
    1. Collection of Seed

    1. Seed Sampling Strategy, sample size and seed health
    Regeneration
    2. Seed germination and seedling raising

    3. Seed harvesting
    4. Drying
    5. Packaging

    2. Germination and vigure testing, dormancy breaking, stratification and vernalization treatment etc.
    3. Number of times, seed collection and extracting techniques
    4. drying methods, slow drying and moisture control testing
    5. Hermetic versus nonhermetic, container types, equipment and packaging environment
    Conservation
    6. base and active collection
    7. Viability monitoring

    8. Seed for regeneration

    6. Seed storage
    7. Sampling, number of seeds, methods(for viability and vigour) and monitoring interval
    8. Threshold viability and sampling
    Characterization and evaluation
    9. Seed germination
    10. Seedling transplanting

    9. As in(2)
    10. suitable site collection
    Distribution
    11. Withdrawing sample and packaging
    12. Documentation

    11. Protecting against temperature and moisture
    12. Barcode versus manual

    Seed Germplasm Collection
    Proper seed sampling techniques have to be applied when collecting seeds from the natural/planted forests. Fully ripened seeds with high initial viability retain it longer than immature seeds (Harrington, 1970; Stein et al., 1974). Knowledge of the right stage and time of maturity is essential for collection of abundant quantity of healthy and vigorous seeds at an economic cost. Physical and biochemical changes occurring during maturation of seeds may determine the proper time of seed collection, storage and testing methods.

    Seed Packaging and Storage
    Seed for long-term storage at –18oC must be hermetically sealed at low seed moisture content of 8 to 12 per cent for most forestry species. Seed with high moisture content in subzero temperatures suffer from freezing injury. Metal can or aluminum-polyethylene pouches can be used for storing the seeds.

    Germplasm Storage
    Conventional seed storage is a relatively cheap method of conservation of wide range of germplasm. According to Harrington (1972), among all the ex situ conservation strategies the easiest and least expensive method of preserving the world’s existing plant genotypes would be conventional seed storage. Seed storage plays a complementary role in germplasm conservation, as temporary means until ex situ stands are established as a safety measure against disastrous losses for limited number of seed lots (Phartyal et al., 2002). So far, only the orthodox seeds (desiccation and chilling tolerant like most of the genera of Fabaceae) have been stored successfully in the genebanks. Generally, knowledge about seed physiology of most forest species is scarce or non-existent. Royal Botanic Garden, Kew, UK with the aim of working in the field of conservation of forest genetic resources started the Millennium Seed Bank Project. It aims to collect and conserve seed of 10 per cent of the world’s flowering plants, mainly from within the drylands, through international effort, by the year 2010. Under this project, a unique seed database has also been developed for information on seed storage behaviour (Tweddle et al., 2002).

    Seed Germination, Dormancy and Pretreatments
    Germinating the seed under optimum conditions is the standard test for assessing the viability of the seeds. Temperature, media and light are the critical factors affecting the seed germination. Optimum temperature required for germination varies with ecotypes within a species also. Determination of cardinal temperature and thermal time for seed germination rate would facilitate the seed bank personnel to select suitable sowing season and agro-climatic zone for introduction of species in field for regeneration and as an in situ conservation stand (Phartyal et al., 2002). Some of the woody species, especially those from tropical highlands and the temperate zone produce seeds, which in spite of being viable fail to germinate promptly even under the optimum germination conditions, this is due to a state called dormancy. Dormancy has evolved as a strategy to avoid germination under conditions where seedling survival is likely to be low. Dormancy has been classified as physical, chemical, mechanical, physiological, morphological and morpho-physiological (Baskin and Baskin, 1998).The most commonly used pre-treatments are soaking seeds in cold and hot water, chemical or mechanical scarification, warm and cold moist stratification, etc. Thus, adequate knowledge about optimum germination, dormancy type in the seed and pretreatments for overcoming it are essential for viability test of the seed to produce uniform results for regenerating the species during the course of its long term preservation.

    Seed Viability Monitoring
    A good seed storage facility will not improve the seed lot quality, it only helps to slow down its deterioration rate. Seed lot storage, therefore, have to be monitored at set interval so that seed lots deteriorated below the viability threshold limit could be withdrawn in time for regeneration to prevent genetic erosion in storage. Monitoring interval should be at every three to five years in the first fifteen years for lots with high initial seed quality in long term storage, followed by more frequent interval after that. The ideal storage conditions are always those, which reduce the growth processes of respiration and transpiration to the lowest possible degree, without impairing the inherent vitality and strength of the seed embryo. The main shortfalls of the current system of assessment of viability is that the seed is consumed in germination tests and that it is time-consuming and costly to do. Ellis et al. (1980) proposed a sequential seed sampling method to cut down seed used, but it has not been adopted because of the increasing labour involved in implementing the method. Presently, research efforts are concentrated on finding non-destructive ways to evaluate seed viability in storage.

    Seed Vigour and Viability Testing
    Where germination tests for some reason (deep dormant seeds or extremely slow germinating seeds) are inconvenient or unreliable, germination potential may be tested by indirect methods viz. viability tests (Schmidt, 2000). Viability tests indicate the potential germinability of seeds, which can reflect expected germination in the nursery. Some of the quick routine indirect tests of viability are the triphenyl tetrazolium chloride (TTZ), excised embryo test, X-radiography and cutting test.

    The main limitation of the germination test is its inability to detect quality differences among seed lots with high germination percentages. So, in such cases vigour tests are more sensitive for detecting the differences. Vigour tests are very important for evaluation of seed quality of stored seeds in a germplasm bank. Decline in vigour is quite difficult to measure, yet several vigour tests have been developed for use for various purposes. The tests include electrical conductivity of seed leachates, accelerated ageing test, excised embryo test, cold test, germination speed, germination value, germination time, germination index, etc. These vigour tests would be very important for assessing the decline in physiological quality of seeds even if their viability remains unchanged during prolonged conservation.

    Conclusion

    So far, gene-banking consists of species with orthodox seeds. Plant genetic resource conservation delves into other forest, types especially tropical rain forests which consist of many rare, endangered and valuable species that produce recalcitrant seeds, for them the problems and techniques would be different than the ones discussed above. Today, inspite of large scale use of forestry species for diverse uses, adequate knowledge on their seed technology and regeneration is deficient. For seed gene banks, seed research is more a science than technology. The storage behaviour of forestry species needs to be thoroughly investigated to develop strategies for their conservation as seeds in gene banks. Quick, low-cost, accurate, non-destructive seed vigour and seed viability tests for predicting seed quality during viability monitoring are necessary. The use of ultra-dried seed storage and ultra-low-temperature cryopreservation techniques are under experimentation and once developed would be effective tools for conservation of our valuable forestry germplasm.

    References

  • ABC Science. 2008. Global seed bank to be Noah’s Ark of food. [Available at: http://www.abc.net.au/science/articles/2008/02/25/2171649.htm]

  • Baskin, C.C. and Baskin, J.M. 1998. Seeds ecology, biogeography and evolution of dormancy and germination. San Diego, Academic Press. 666 p.

  • Ellis, R.H.; Roberts, E.H. and Whitehead, J. 1980. A new, more economic and accurate approach to monitoring the viability of accessions during storage in seed banks. Plant Genetic Resources Newsletter, 41: 3-18.

  • Engelmann, F. and Takagi, H. 2000. Cryopreservation of tropical plant germplasm: Current research progress and application. Tsukuba, Japan International Research Centre for Agricultural Sciences. 496 p.

  • Food and Agriculture Organization of the United Nations. 1997. The new revised text of the International Plant Protection Convention 1997. Rome, F.A.O.

  • Food and Agriculture Organization of the United Nations. 1998. The state of the world’s plant genetic resources for food and agriculture. Rome, F.A.O.

  • Food and Agriculture Organization of the United Nations/Inernational Plant Genetic Resources Institute. 1994. Genebank standards. Rome, FAO/IPGRI. 46 p.

  • Harrington, J.F. 1970. Seed and pollen storage for conservation of plant gene resource. In: Genetic Resources in plants, their exploration and conservation. London, International Biological Programme.

  • Harrington, J.F. 1972. Seed storage and longevity. In: Kozlowski, T.T. Ed. Seed biology V.3. New York, Academic Press. pp. 145-245.

  • Phartyal, S.S.; Thapliyal, R.C.; Koedam, Nico and Godefroid, S. 2002. Ex situ conservation of rare and valuable forest tree species through seed-gene bank. Current Science, 83(11): 1351-1357.

  • Roche, L. 1975. Report on a pilot study on the methodology of conservation of forest genetic resources. Rome, F.A.O.

  • Schmidt, L. 2000. Guide to handling of tropical and subtropical forest seed. Humlebaek, Danida Forest Seed Centre. 511p.

  • Stein, W.I.; Slabaugh, P.E. and Plummer, A.P. 1974. Harvest, Processing and storage of fruits and seeds. In: United States Department of Agriculture. Forest Service. Seed of woody plants in the United States. Washington DC., USDA Forest Service. pp. 300-320.

  • Tay, D.C.S. 2007. Seed Technology in plant germplasm conservation. In: Basra, Amarjit S. Ed. Handbook of seed science and technology. Jodhpur, Scientific Publishers. pp. 731- 748.

  • Tweddle, J.C.; Turner, R.M. and Dickie, J.B. 2002. Seed information database. [Available at: http://www.rbgkew.org.uk/data/sid]

  •  India’s National Action Plan on Climate Change

    Prime Minister Dr. Manmohan Singh on June 30, 2008 released India’s national action plan on climate change in New Delhi, prepared under the guidance and direction of the prime minister’s council on climate change. Members of the prime minister’s council on climate change, senior members of the union cabinet, environmental activists and senior government officials, were present on the occasion.

    Speaking on the occasion, Dr. Singh said that the release of the plan reflected the importance the government has attached to mobilising its energies to meet the challenge of climate change.

    The national action plan focuses attention of eight ‘priority national missions’:
    1. Solar energy
    2. Enhanced energy efficiency
    3. Sustainable habitat
    4. Conserving water
    5. Sustaining the Himalayan ecosystem
    6. A “green India”
    7. Sustainable agriculture
    8. Strategic knowledge platform for climate change

    The national mission on solar energy occupies a pre-eminent place, whose success, the prime minister said, has the potential of transforming the face of India.

    Dr. Singh added that India believed that every citizen of this planet should have an equal share of the planetary atmospheric space and therefore, long-term convergence of per capita GHG emissions was the only equitable basis for a global agreement to tackle climate change. In this context, the Prime Minister reaffirmed India’s pledge that as it pursued sustainable development, its per capita GHC emissions would not exceed the per capita GHG emissions of developed countries, despite our developmental imperatives.


    SEED SOURCE VARIATION IN SOME IMPORTANT NATIVE TREE SPECIES OF GHARWAL HMALAYA

    N.P. Todaria and Bhupendra Singh
    Department of Forestry, H.N.B. Garhwal University, Srinagar - 264 174


    Introduction

    F orest species with tremendous amount of heterozygosity also exhibit considerable variation in seed traits like hard seed coatedness, dormancy, pretreatment requirement, length of stratification period, morphological variation in seed characters, germination rate and cardinal temperature for optimum germination and storability in addition to variation in economic characteristics like bole straightness, volume, disease and pest resistance. The knowledge of genetic variation within a species is a prerequisite for developing effective tree improvement programme, which begins with the scanning of available variations in the entire range of species distribution and delimitation of provenances/seed sources capable of providing best adapted trees (Suri, 1984). Provenance tests in native species are desirable to screen the available variation for higher productivity and future breeding work (Burley and Wood, 1976). Selection of best provenance of desired species for a given site or region is necessary for achieving maximum productivity both in plantation forestry and agroforestry systems (Subramanium et al., 1992).

    Tree species whose distribution extends to a large geographical area with varying climate and topography are usually known to have local populations adapted to their environment. The degree of adaptations vary with relative rates of environmental stress or sometimes, sudden change in topography, aspect and soil, resulting in the corresponding genetic variation in the species. These variations within the population of tree species have been exploited in the selection of superior provenance for a given site and for evolving strategies for conservation of genetic diversity within population of trees species (Thapliyal and Dhiman, 1997). Department of Forestry, H.N.B. Garhwal University, has been carrying out seed source studies on forest species distributed with in western Himalaya. The three region of western Himalaya i.e. Garhwal, Kumaun and Himachal Pradesh show a great degree of environmental variation and thus give us a opportunity to carry out such studies. The work carried out during last 12 years has been summarised herein.

    Seed Source Variation in Seed Size and Seed Weight

    Variations obtained for seed morphology has been assigned to seed source. Variation and inheritance of seed size among seed source have been documented for different native tree species of Garhwal Himalaya, including Acacia catechu (Todaria et al., 2004); Albizia chinensis (Dhanai et al., 2003); Grewia oppositifolia (Uniyal, 1998); Quercus leucotrichophora (Saklani, 1999) Celtis australis (Singh et al., 2006) and Terminalia species (Chauhan et al., 2007). Seeds of Acacia catechu collected from different seed sources provided significant variation in seed length, breadth, thickness and weight among most of the seed source (Todaria et al., 2004). Dhanai et al. (2003) collected 13 provenances of Albizia chinensis in Western Himalaya (Uttarakhand and Himachal Pradesh) and found that seed morphology of A. chinensis significantly varied among the provenance but altitude of seed collection was non-effective.

    Seed weight is mainly influenced by maternal factors and is under genetic control (Dhanai et al., 2003). In Celtis australis higher elevational seed sources produce heavier seed and seedlings (Singh et al., 2006). Relative high seed weight is often desirable since it is often correlated with rapid germination and good seedling establishment (Uniyal et al., 2003, Singh et al., 2006). Seed weight of Quercus floribunda was significantly inversely correlated with altitude of seed origin. This is an indication that some of these phenotypic variations may also have genetic base when species distribution is wide (Uniyal et al., 2002). Altitude did not affect much of the characters in species having short altitudinal gradient of distribution like Quercus spp. The seed weight of Pinus roxburghii was found most variable character as compared to other seed characters and it may be the most heritable character (Todaria et al., 2003) (Table 1).

    Table 1. Morphological variation of some of the native tree species (value in parenthesis indicated range between seed source)

    Species Pod Lenght
    (cm)
    Pod Width
    (cm)
    Seed Lenght
    (cm)
    Seed Width
    (cm)
    Seed Weight
    (gms/100 seeds)
    Acacia catechu 7.37
    (6.16-8.13)
    1.21
    (1.02- 1.50)
    6.68
    (5.17-7.62)
    5.69
    (4.45-6.32)
    3.87
    (2.23-5.17)
    Albizia chinensis 10.66
    (8.92-13.31)
    1.57
    (1.38-1.87)
    6.14
    (5.79-6.53)
    4.52
    (4.19-5.08)
    2.19
    (1.79-2.66)
    Albizia lebbeck 19.79
    (11.15-24.49)
    2.84
    (2.4-3.39)
    9.00
    (0.75-1.17)
    7.42
    (0.57-0.81)
    6.75
    (4.43-9.89)
    Albizia procera 12.47
    (11.22-14.58)
    1.56
    (1.28-1.76)
    5.60
    (5.00-5.90)
    4.50
    (4.2-5.0)
    4.24
    (3.29-4.59)
    Celtis australis - - 5.40
    (4.50-6.30)
    4.40
    (4.0-4.70)
    0.67
    (0.48-0.83)
    Dalbergia sissco 6.11
    (4.88-8.05)
    0.94
    (0.58-1.12)
    8.20
    (7.6-9.8)
    4.6
    (4.1-5.3)
    1.70
    (1.35-2.23)
    Grewia optiva - - 6.77
    (6.3-7.7)
    5.78
    (4.6-6.8)
    10.61
    (7.04-14.32)
    Pinus roxburghii - - 11.10
    (8.8-12.7)
    5.90
    (5.4-6.6)
    115.9
    (68.9-155.0)
    Quercus glauca - - 19.6
    (10.3-23.0)
    12.2
    (10.4-13.9)
    139.79
    (106.6-183.3)
    Quercus leucotrichophora - - 21.5
    (18.9-24.1)
    11.4
    (9.8-12.5)
    151.6
    (138-188)
    Quercus floribunda - - 32.40
    (27.6-3.66)
    14.99
    (13.95-16.39)
    305.29
    (201.0-363.0)
    Terminalia bellirica - - 34.00
    (2.9-4.0)
    7.50
    (6.2-10.2)
    910.00
    (546-1840)
    Terminalia chebula - - 4.60
    (4.0-5.3)
    7.90
    (6.6-9.6)
    1321.0
    (921-2.60)
    Terminalia tomentosa - - 4.10
    (3.0-4.1)
    8.90
    (8.7-9.4)
    125.00
    (96-179)

    In Pinus brutia, almost all the seed characteristics studied by Isik (1986) showed positive correlation with rainfall, longitude and latitude of seed origin. A significant relation was found between seed weight and growth of seedlings in nursery and plantation, indicating it to be an important trait for early selection of seed source. Bagchi and Dobriyal (1992) reported seed source variability in seed and seedling traits of some Indian provenances of Acacia nilotica. Trends of germination and seedling growth rates with seed size may stabilize seed size variation within a species (Burslem and Miller, 2001). Seed size has been considered to be one of the phenotypically least flexible characters in many tree speices (Fenner, 1992), since individual seeds within bulk sample can show substantial variation in size (Stamp, 1990; Counts and Lee, 1991; Banzonetz and Scheiner, 1994). Environmental factors are also known to influence both germination and seed size (Fenner, 1992; Wolfe, 1995). Variation in seed size is probably due to different positions on the mother plant, resulting in differences in seed filling (Gutterman, 1992).

    Seed Source and Seed Germination

    Studies on some forest tree species also revealed that germination rate and germination period may change from provenance to provenance and it may be due to the variation in seed size and weight among the seed source (Chauhan, 1998; Uniyal, 1998; Saklani, 1999; Dhanai, 2003; Singh, 2004). Seed size and weight influence seed germination behaviour and seedling development. Heavy and large seeds of Terminalia bellerica and Acer oblongum performed better in terms of germination pattern while T. tomentosa and T. bellerica gave better results in term of seedling development after 6 months growth (Negi and Todaria, 1997). Another study (Khan et. al., 1999) revealed that large or 1-seeded fruits of Mesua ferrea gave better germination in laboratory and greenhouse. Further, seedling survival and vigour were greater for seed from large rather than small fruits. Studies relating seed source variations resulting in germination differences among populations have been recorded in Celtis australis (Singh et. al., 2004), Albizia lebbeck (Todaria et. al., 2002), Albizia chinensis (Dahani, 2003), Grewia optiva (Uniyal et. al., 2003), Quercus leucotrichophora (Saklani et. al., 1999) and Terminalia species (Chauhan et. al., 2007).

    Wide variation in seed germination, seedling height and diameter are essentially genetic in origin. These have been confirmed in studies of some native species of Garhwal Himalaya such as Terminalia tomentosa, T. bellirica, T. chebula (Chauhan et. al., 2002), Grewia optiva (Uniyal et. al., 2003), Acacia catechu (Todaria et. al., 2004), Albizia lebbeck (Todaria et. al., 2002), Albizia chenensis (Dhanai, 2003) and Celtis australis (Singh et. al., 2004)Seed germination decreased as the altitude of seed collection increased in Acer oblongum, Anogeissus latifolia, Sapindus mukorossi, Kydia calycina and Toona ciliata (Todaria and Negi, 1995). Chauhan et. al. (1996).also concluded that seed germination of Alnus nepalensis was strongly influenced by the seed source (altitude), the seed germination of A. nepalensis decreased with increasing altitude of seed collection. The germination physiology is adapted to a particular environment as the requirement of certain range of temperature is also involved in determining the plant distribution. This was again proved when some non-germinated seeds of Acer oblongum, Anogeissus latifolia, Kydia calycina, Sapindus mukorossi, Terminalia bellirica and T. chebula transferred from ower to higher temperature regime, resulted in higher germination percentage (Negi and Todaria, 1993). However, seed germination in Quercus floribunda was significantly positively correlated with seed source at all the tested temperature treatments (Saklani et. al., 2001). Variation in germination of Quercus floribunda may be due to genetic differences, however, environmental factors may also play a role in changing the component of seed size and weight as they influence the seed germination.

    The germination of seeds is strongly influenced by variation in temperature, water stress and light requirements. These factors often show significant interaction in their effect on germination (Singh et al., 2007). The magnitude of seed germination response in Q. floribunda varied among seed population, however, 25oC constant temperature induced optimum germination in all the seed population followed by 20oC temperature. But there was very low average germination at 30oC temperature (Saklani et al., 2001). The seed germination of Terminalia species also varied between seed sources. In Terminalia species optimum temperature for seed germination was 25oC constant and 30/25oC alternate temperatures. In Albizia lebbeck also 25oC constant and 30/20oC alternate temperature produced optimum germination irrespective of seed source (Todaria et al., 2002). Singh et al. (2004) tested different seed sources of Celtis australis at different constant and alternating temperatures and found that seed germination in C. australis was strongly influenced by seed source and temperature regimes. However, the optimum seed germination was obtained at 25oC irrespective of seed sources.

    The optimal germination temperature for Momordica charantia L. seeds is between 25oC and 28oC, and poor germination is common at sub-optimal temperatures (Peter et al., 1998; Lin and Sung, 2001). Alternating temperatures were the best option for the germination of Trema micrantha seeds, as the seeds of Trema generally do not germinate at constant temperature (Castellani and Aguiar, 2001). Germination physiology generally indicates temperature as a range, within which the optimum is mentioned as the temperature at which the highest percentage of germination occurs within the shorter period of time. Further, each species has own temperature requirement for germination, suggesting a possible genetic control (Khullar et al., 1991; Todaria and Negi, 1995).

    Seed sources also significantly influence seedling growth. Several studies have been carried out to examine the variation in seedling traits among the seed sources in different species in Garhwal Himalaya viz. Terminalia tomentosa, T. bellerica T. chebula (Chauhan, 1998); Grewia optiva (Uniyal et. al., 2003); Quercus leucotrichophora (Saklani, 1998); Albizia chinesis (Dhanai, et al., 2003); Celtis australis (Singh et al., 2006). Significant variations have been recorded for all the growth traits among the 13 provenances of A. chinensis. The growth performance of Celtis australis seedling was positively significantly correlated with elevational range of seed sources. The significant correlation was also recorded between seed weight and seedling growth of C. australis (Singh et al., 2006). Seedling growth of Grewia optiva also significantly varied among the seed sources (Uniyal et al., 2003). Uniyal et al. (2007) also reported that heavier seeds of Azadirachta indica produce larger seedling as compared to the lighter ones.

    Conclusion

    Selection of suitable seed source is key for producing quality planting stock. The quality of seed and seed weight are the most variable characters and are directly correlated with the germination and growth of the seedlings. The range of temperature requirement for optimum germination has implication for nursery germination. Thus, seeds of particular species should be sown in nursery as and when specific range of temperature is available.

    References

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  • Saklani, K.P. 1999. Altitudinal and seasonal variation in relation to fodder quality of oak (Quercus leucotrichophora A. camus ex. Bahadur) in Garhwal Himalaya. D. Phil. theses, H.N.B. Garhwal University, Srinagar. 111p.

  • Saklani, K.P.; Chauhan, S.; Bhatt, B.P. and Todaria, N.P. 2002. Effect of seed source and temperature on seed germination of Quercus floribunda Lindley Ex. Rehder in Garhwal Himalaya. Journal of Plant Biology, 28(3): 307-310.

  • Singh, Bhupendra; Bhatt, B.P. and Prasad, P. 2004. Effect of seed source and temperature on seed germination of C. australis L.: A promising agroforestry tree-crop of central Himalaya, India. Forests, Trees and Livelihoods, 14(1): 53-60.

  • Singh, Bhupendra; Bhatt, B.P. and Prasad, P. 2006. Variation in seed and seedling traits of Celtis australis, a multipurpose tree, in central Himalaya, India. Agroforestry Systems, 67(2): 115-122.

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  • Todaria, N.P.; Bagwari, H.K. and Chauhan, D.S. 2004. Effect of seed source, temperature and light on seed germination of Acacia catechu. Indian Journal of Tropical Biodiversity, 12: 43-47.

  • Todaria, N.P.; Chauhan, Shashi and Sachan, M.S. 2003. Variability studies in Albizia lebbek from Garhwal Himalaya. Range Management and Agroforestry, 24(1): 35-37.

  • Todaria, N.P. and Negi, A.K. 1995. Effect of elevation and temperature on seed germination of some Himalayan tree species. Plant Physiology and Biochemistry, 22: 178-182.

  • Todaria, N.P.; Ram Kumar and Chauhan, Shashi. 2003. Effects of temperature and light on seed germination of Pinus roxburghii. Journal of Tree Sciences, 22(1-2): 69-72.

  • Todaria, N.P.; Saklani, K.P. and Sachan, Manoj. 2004. Variation in pod and seed characteristics of Acacia catechu collected from Garhwal Himalaya. Indian Forester, 130(1): 53-61.

  • Todaria, N.P.; Singh, Bhupendra and Ram Kumar. 2002. Seed source variation and germination studies in Albizia lebbek from Garhwal Himalaya. Journal of Tree Science, 21(1-2): 27-34.

  • Todaria, N.P.; Uniyal, A.K. and Bagwari, H.K. 2003. Source variation in cone and seed characteristics of Pinus roxburghii in Garhwal Himalaya. Annals of Forestry, 11(1): 8-19.

  • Uniyal, A.K. 1998. Provenance variation in seed and seeding of Grewia optiva Drumm. D. Phil. thesis, H.N.B. Garhwal University, Srinagar. 143 p.

  • Uniyal, A.K.; Bhatt, B.P. and Todaria, N.P. 2002. Provenance variation in seed characteristics of Grewia oppositifolia: A promising agroforestry tree-crop of central Himalaya, India. Indian Journal of Forestry, 25(2): 209-214.

  • Uniyal, A.K.; Bhatt, B.P. and Todaria, N.P. 2003. Effect of provenance variation on seed and seedling characteristics of Grewia oppositifolia: A promising agroforestry tree-crop of central Himalaya, India. Plant Genetic Resources Newsletter, 136: 47-53.

  • Uniyal, A.K.; Singh, Bhupendra and Todaria, N.P. 2007. Effect of seed size, sowing orientation and depth on germination and seedling growth in neem, Azadirachta indica. Seed Technology, 29(1): 68-75.

  • Wolfe, L.M. 1995. The genetics and ecology of seed size variation in a biennial plant Hydrophyllum apendiculatum. Oecologia, 101: 343-352.


    ENVIRONMENTAL
    UPS and DOWNS


    Environmental Ups

    With the aim of creating awareness among different target groups about the consequences of loss of biodiversity and how we contribute to conserve this invaluable wealth, the United Nations has proclaimed 22nd May, the International Day for Biological Diversity (IBD).

    In yet another attempt to clean the Yamuna, interceptor sewers will be installed which will complete the cleaning of the river before the Commonwealth Games in 2010 – says Delhi Chief Minister Sheila Dikshit.

    Riding high on its achievements, the Delhi Metro has added another feather in its cap when it became the first railway project in the world to be registered by the UN Framework Convention on Climate Change. This feat would be of tremendous financial help in the name of earning carbon credits, which is measured in terms of reduction of carbon dioxide by adequate measures.

    In spite of mounting developmental pressures, the hill states of the country including Uttarakhand continue to hold their forest cover, registering miniscule losses, revealed in the latest ‘State of Forest Report’, released by Forest Survey of India.

    Scientists at G.B. Pant Institute of Himalayan Environment and Development, Almora reported that the Gangotri glacier has retreated at much lower rate between 2004 and 2005. This is claimed to be the first and accurate report on glacier retreat measurement using high precision GPS in India.

    There is something positive for wildlife enthusiasts. The population of elephants in the wild has jumped by more than a thousand. As per the latest 2007 survey, it stands between 18,663 and 18,713. This is excluding the figures from the north-eastern states for which the data are yet to be compiled.

    If two scientists at Los Alamos National Laboratory are correct, people will still be driving gasoline-powered cars 50 years from now, churning out heat-trapping carbon dioxide into the atmosphere – and yet that carbon dioxide will not contribute to global warming.

    Prime Minister Dr. Manmohan Singh said in the Rajya Sabha that the government was preparing a National Action Plan at the highest level to deal with the issues of climate change, including the remedial measures for mitigating the impact on food grains production.

    Climate scientists at Kathmandu-based International Centre for Integrated Mountain Development (ICIMOD) have mooted a comprehensive regional approach to fight the looming human-induced climate crisis and avert future social conflicts over dwindling of natural resources, especially water.

    The Ministry of Environment and Forests, Govt. of India plans to take up plantation in over six million hectare of degraded forests of the country in two phases in the next 10 years at an estimated rate of around one lakh trees per hectare of land. This programme will be funded out of the money collected under Compensatory Afforestation Fund Management and Planning Authority.

    Fourteen tiger cubs have been regularly spotted in Ranthambore National Park in western Rajasthan, which is a rare piece of good news in the country’s fight to protect its dwindling population of big cats from poachers and habitat destruction.

    The Government of India is planning to compile detailed information on Indian forests to understand the health of the green cover in the country, which will also help quantify contribution of forests in terms of monetary and realistically measurable value.

    The Ministry of Environment and Forests, Government of India has decided to create four new tiger reserves one each in the states of Madhya Pradesh, Maharashtra, Orissa and Uttar Pradesh; and to upgrade Nagarohole National Park as a separate reserve, which presently is a part of Bandipur Tiger Reserve.

    A satellite-linked fire alert system called Fire Alert and Message System (FAMS) developed by NASA and the University of Maryland, US, currently on trial in Madhya Pradesh, is turning out to be an effective tool in saving wildlife and biodiversity from forest fires.

    Union Minister of State for Forests and Wildlife S. Regupathy said that forest cover in India would reach 30 per cent of the total geographical area by 2012.

    Environmental Downs

    Studies conducted by the New Delhi based Indian Agricultural Research Institute have pointed to a possible loss of 4 to 5 million tonnes in the overall wheat production with every one degree centigrade increase in temperature throughout the growing period of the crop.

    Environmental pollution may trigger diabetes, scientists suspect in the US. In the commentary published in the recent edition of the Lacet, Cambridge scientists, Oliver Jones and Julian Griffin highlighted the need to research the possible link between certain pollutants and diabetes.

    The United Nations Environment Programme’s latest report entitled ‘In Dead Water’ said that climate change might slow down the global flow of ocean currents, which flush and clean the continental shelves and are critical to maintaining water quality, nutrient cycling and the life-cycle patterns of fish and other marine life in more than 75 per cent of the world’s fishing grounds, which would eventually adversely impact the fish production globally.

    Around 728 sq km of forests – roughly half the size of Delhi – have been wiped out in two years in India – reports ‘State of Forest Report’, released by the Forest Survey of India.

    An ecological disaster sparked by global warming is waiting to strike the remote Arctic tundra, warns a study funded by the National Science Foundation of USA. According to the study, the high latitude tundra and boreal forest ecosystems comprise a significant 30 per cent of the planet’s total soil carbon, much of which is locked in permafrost. A warming climate could cause the permafrost to melt and release its carbon stores into the atmosphere where it would contribute greatly to the greenhouse effect.

    The water supplies will drastically shrink as snow disappears and ice melts. The lack of safe water will most probably trigger outbreaks of diarrhoea and other diseases, says a report of the World Health Organisation.

    The Uttar Pradesh Pollution Control Board and Ganga Research Laboratory at the Institute of Technology, Banaras Hindu University noticed that pollution level in the Ganga had reached alarming proportions, and Ganga water in Varanasi was not fit even for bathing and agricultural purposes.

    The groundwater reserves are depleting at an alarming rate in Punjab besides a serious decline in the quality of water at many places due to overuse of the available resources, says a study conducted by Water Resources and Environment Directorate of Punjab Irrigation Department.

    The vulture may soon become extinct in India if their population continues to decline at the present rate despite measures taken by the government as 99 per cent population of this bird has declined at the rate of around 50 per cent every year, states Bombay Natural History Society.

    The world’s wildlife population reduced around a quarter since the 1970s, states World Wide Fund for Nature.

    The U.N. Environment Programme has reported sharp decline in the number of migratory birds, which are true citizens of the world, appears to be losing out to developmental pressures.

    The research by 15 scientists from 13 different countries found that overfishing is largely to blame for putting 11 species of oceanic sharks rays out of 21 species under study, to nearly wiped out, says IUCN’s Shark Specialist Group (SSG).

    The volume of water that flows in the river Ganga through the ancient city of Kashi (now Varanasi) has decreased with water level dropping to all time low this summer. According to Prof. U.K. Chaudhary of B.H.U., Varanasi “The water level of Ganga in Varanasi had dropped by 1.5 to 2 metre over the past two decades”.

    The Himalayan quail again finds mention in the critically endangered category of the 2008 Red Data List of IUCN. The bird has been in critically endangered category list since 1994.

    Mass extinction of plants and animals could have a severe impact on the living standards of the poorest people on the planet and cost up to Ł40 billion a year, states ‘The Economics of Ecosystems and Biodiversity Review’ which analysed the financial impact of the loss of natural life.


    SIGNIFICANCE OF SEED PATHOLOGY IN FOREST TREES

    K. Vishunavat and J. Kumar
    Department of Plant Pathology, G.B. Pant University of Agriculture and Technology,
    Pantnagar- 263 145


    Introduction

    F orest covers about 30 per cent of the Earth land area (Food and Agriculture Organisation of the United Nations, 2000). Tropical and subtropical forest comprise 56 per cent of the world forest while temperate forest accounts for 44 per cent. Forest plantations make up only 5 per cent of all forest; rest is natural forest. Covering about one third of the Earth’s land surface, forests provide many benefits. The most notable benefits are an estimated 5,000 commercial products, such as lumber, paper, turpentine, and others, worth billions of dollars a year. Forests also provide refuge from hectic urban life and opportunities for many forms of recreation. In many poorer nations, forests are a source of wood for cooking and heating.  Forests are also home to many of the world’s species.  Forests benefit us indirectly by protecting watersheds from soil erosion, and keep rivers and reservoirs relatively free of silt.  Forests reduce the severity of floods and facilitate aquifer recharge which assists in the recycling of water, oxygen, nitrogen, carbon and other nutrients. Despite great benefits of forests, only 13 per cent of the world’s forestland is under any kind of management. In addition, only 2 per cent of the world’s forests are protected in forest reserves.

    Since the advent of agriculture, about 33 per cent of the world’s forests have been cleared and converted to other uses, mostly farms and human settlements.  To date, the United States and Africa have both lost about one-third of their forests, while Brazil, the Philippines and Europe have lost 40 per cent, 50 per cent and 70 per cent, respectively.  Moreover, deforestation continues virtually everywhere.  By one estimate, 17 million hectares of tropical rain forests are leveled each year. The World Bank estimates that within a decade the number of tropical countries that export wood will drop from 33 to about 10. In India, forestland is shrinking by 1 million hectors per year. At its current rate of harvest, China will lose all of its commercial forests within ten years. In the 1980s, softwood harvest on the West Coast of the United States exceeded sustainable yield by 25 per cent on privately owned land and 61 per cent on national forests. In developing countries, for every 10 trees cut down only 1 tree is replanted. In Africa, the ratio is 29 to 1. Replanting programs need to be prioritized by those benefiting wildlife and species diversity verses those having lesser benefit.

    The ever-increasing impact of human activities on the environment makes the conservation of natural resources, including biological diversity, an urgent and critical task leading to changes and introduction in the flora and fauna in forest with the new pathogens or virulent races of the pathogens leading to newer diseases, which may be severe and thus lead to substantial losses.

    Importance of Forest Pathology

    The science of Forest Pathology as profession began when a Germen forester Robert Hartig, fathered forest pathology as a science in the 1870s. Forest pathology is the branch of plant pathology that deals with diseases of woody plant growing in natural forests, in plantation, in urban environment and microbiological deterioration of forest product after harvests. Danida Forest Seed was established in 1968 as Danish / FAO Forest Tree Seed Centre. The name was subsequently changed to Danida Forest Seed Centre (DFSC) in 1981. The main objectives of DFSC are to promote supply, production and use of genetically and physiologically improved seed material for tree planting programme in forestry and agro-forestry in developing countries.

    Diseases of forests are caused by fungi, bacteria, nematodes, seed plants, viruses,mycoplasmas, as well as unfavorable environmental factors including heat, cold, drought, flooding, air pollution, nutrient deficiencies, and adverse physical factors or toxic substances in soil. Devastating losses due to diseases such as Dutch elm disease, Chestnut blight, pine blister rust, oak wilt, Cyprus canker, etc., in trees have been experienced from time to time.

    Deterioration of tree seeds by fungi involves problems that differ in many aspects from those of grains. For example, tree seeds are exposed to many conditions before storage that permit the development of mold fungi. Hence, it is important to know the characteristics of fungi associated with important tree species, what damage they cause, where and when and under what circumstances the damage occurs, and what could be done to prevent the damage.

    The science of tree seed pathology is still very young. The occurrence and distribution of most of the tree seed pathogens is well realized but there is very little understanding of their impact on seed production, seed quality and seed viability. Interest in these problems has been growing steadily in the past few decades, but recently this interest has changed in to serious concern mostly because of the problems encountered in the renewal and management of the forests, which are a major resource worldwide.

    Since the inception of the forest pathology, pioneering studies have been carried out on many disease problems of seeds, seedlings in nurseries, plantations and natural forests and their management. Significant contributions were made on the aspects of timber pathology and mycorrhizae. More than 160 fungi causing diseases and decays of both hardwood and soft wood species were studied. Seeds of mandated tree species were screened for seed borne pathogens and their management was evolved to minimize losses in storage and nursery. In Dalbergia sissoo among the the nursery diseases, the most notable ones are damping-off, and Rhizoctonia leaf and web blight (Mehrotra, 1998). The Melampsora rust (Pei and McCracken, 2005), Uredo sissoo, Maravalia achroa and Maravalia pterocarpi have also been reported to cause foliage infections in nursery seedlings. A Meliola sp. causes a sooty mould of seedlings. Certain seedling diseases like Rosellina root rot poplar scab (also known as ‘Summer Leaf Drop’) caused by a fungus Venturia populina (anamorph Pollaccia elegans), Bipolaris leaf blight diseases of poplars, Cylindrocladium leaf and twig blight of eucalypts and Pseudocercospora needle blight of pines have been investigated and management have been worked out extensively.

    In plantations, some of the important diseases include Trichosporium wilt of Casuarina equisetifolia, Fusarium wilt of Dalbergia sissoo, pink disease of Eucalyptus, Ganoderma root rot of hard wood species, partridge root rot of Shorea robusta caused by Aurificaria shoreae, root rot of Gmelina arborea by Poria rhizomorpha, root rot of Cedrus deodara by Heterobasidion annosum,Armillaria root rot of Abies pindrow, Picea smithiana and Cryptomeria japonica and brown root rot of poplars caused by Phellinus noxius.

    The importance of forests pathology is a function of the economic and aesthetic value of trees and forest products. Disease losses occur virtually in every forest stand in every region of the world. Diseases cause greater growth loss than fire, insects, animals, or weather. Health and vigor of seedlings and their further growth are to a considerable extent dependent on the quality of seeds since seedlings grown from the seeds are the primary source of pathogens. In the past, damping-off fungi, cone rust, a number of cone and seed insects in pines, and fungal damages to acorns (oak), hazelnuts, chestnuts, walnuts and seeds of birch and elm were considered the only major problems in seed and seedling production. Many fungi have now been isolated and studied for their effects on seeds of both conifer and hardwood tree species. Diseases operate in many ways to reduce the yield and quality of timber and other values of trees. Losses begin with seed abortion and deterioration in the germinabilty of seeds and the survival of seedlings. Root pathogen, e.g. Fomes annous and Poria weirii, kill or decrease tree growth. Bark and cambial parasite, e.g. Nectria galligena and Strumella coryneoidea, can girdle and kill trees or produce lesions that lead to decay. Wood discoloring and rotting fungi, e.g. Fomes pini and decrease the merchantability of wood in conifers and hard woods. Foliage diseases, such as Elytroderma deformans needle blight in ponderosa pine and sycamore anthracnose decrease the amenity value of trees, decrease their growth and some times kill them.

    Seed borne infection, causing seed quality losses of Botryodiplodia theobromae on Swietenia macrophylla; Colletotrichum gloeosporoides on Dalbergia cochinensis; Alternaria longissima on Bauhinia sp.; Pestalotiopsis sp. on Cassia bakeriana; Macrophoma sp. on Eucalyptus camaldulensis and Fusarium sp. on Shorea obtuse has been observed.

    Significance of Forest Seed Pathology

    Fungal diseases are serious problems in regeneration and, some times, can cause heavy mortality in nurseries and become established on seedlings. Apart from seed borne fungal pathogens, soil borne pathogens also have been found to cause seedling mortality in forest nurseries. These seedlings being tender in tissues often face difficulty in establishing themselves. Such substandard seedlings when used as planting stock further spread the disease to plantation and forest causing heavy damages.

    Fungi associated with tree seeds vary in different host species, in different regions and in different years (Mittal et al., 1990). Nearly all seeds carry spores of various microscopic fungi either on the surface or within the seed. Superficial mycelium is almost always found because of the ready adhesion of spores to the uneven surface of the seeds. In Norway, spruce and Scotch pine seeds, Sirococcus strobilinus spore load can be as high as 150,000 spores and in some seed lots several hundred thousand spores/gram seed. Under favorable conditions, some spores germinate, the mycelium penetrating into the cotyledons of the seed and feed on the embryo. Several kinds of fungi can be associated with tree seeds. Depending upon their location, the seed-borne fungi can, in general, be classified in two groups: externally seed borne and internally seed-borne. An extensive study on seed borne fungi and their management has been carried out at Danish Forest Seed Centre, Denmark with a view to maintain the quality of tree seeds (Rees and Phillips, 1986).

    Externally seed borne fungi are those that survive on the upper surface of the seeds. Examples are Botryosphaeria, Botrytis, Fusarium, Mucor, Phialophora, Rhizopus, and Trichothecium, etc. They are not usually host specific and may involve more than one host species. A rich microflora has been recorded in Albizia lebbeck seed samples comprising Aspergillus, Chaetomimum, Rhizopus., Penicillum which occurred in a range of 21-48 per cent (Mohanan et al., 2005).

    Internally seed-borne fungi include species of Alternaria, Aspergillus, Botrytis, Botryodiplodia, Caloscypha, Cephalosporium, Fusarium, Phoma, Schizophyllum and Sirococcus. These may cause deterioration of seed quality and pre- or post-emergence mortality of seedlings. Symptoms of seed-borne diseases are usually divided into pre- and post-emergence damping-off. The former consists of reduced emergence and decay of the radicle just emerged from the seed coat; the latter is subdivided into root rot, cotyledon rot, and basal stem rot after the seedlings emerge from the soil. Reduction in seed germination, decay and loss of viability of seeds during storage, and the diseases of seedlings are among the major problems brought by fungal pathogens.

    Losses Due to Seedborne Fungi in Tree Seed and Seedling Diseases

    Damping Off
    Pre- and post-emergence damping off caused by various fungi are the most dangerous diseases affecting conifers as well as hardwood species. Fusarium solani, is a soil inhabitant, in the seeds of Leucaena and Agathis which caused post- emergence damping-off in the nursery and in the out planted seedlings. Fusarium-contaminated seed with species like Fusarium oxysporum, Fusarium equiseti, Fusarium verticillioides, Fusarium incarnatum and Fusarium acuminatum cause high damping-off losses in Pinus ponderosa (Salerno and Lori, 2008).

    Seedling Blight
    Sirococcus blight caused by the seed-borne fungus Sirococcus strobilinus is an important disease of seedlings of several spruce and pine species and of Pseudotsuga. The pathogen attack very young seedlings, killing the primary needles from the base upward. Dead seedlings remain upright and small, black pycnidia usually form at the base of infected needles. Diseased seedlings usually occur randomly, characteristic of seed-borne diseases. Pinus ponderosa show seedlings chlorosis due to seed-borne Fusaria species, like F. oxysporum, F. equiseti, F. incarnatum, F. acuminatum and F. verticillioides. Rhizoctonia solani (Thanatephorus cucumeris) has been found to cause the leaf blight of Cassia fistula, Bauhinia variegata, Dalbergia sissoo and Populus deltoides in the nurseries in western Uttar Pradesh, India. The disease caused blighting and webbing of leaves forming cobweb-like structures. The disease caused premature defoliation and group infection of seedlings due to lateral spread of the disease through contact of the overlapping foliage of the adjoining seedlings (Mahrotra 1998). Leaf spot and leaf blight of Dendrocalamus strictus and due to Cercospora apii and Myrothecium roridum; root rot of Hardwickia binata (Fusarium oxysporium) and leaf blight of Terminalia catappa (Fusarium solani) are important seedling diseases. Myrothecium roridum has also been found to cause leaf spot in Bombax ceiba (Sharma et al., 1985) and Pterocarpus marsupium (Ali and Sharma, 1996). Cercospora dehraduni, a new species from India has been reported (Mishra, 2001).

    Seedling Wilt
    Another important disease transferred by seeds is the Tracheomycosis wilting of plants. This symptom can be elucidated as a reaction of the host to the irritation by the parasite wherein the typical blocking of trachae by thalli, and a yellowish brown, rubber-like substance filling the adjacent parenchymatous cells, are produced. Reduced seedling height and leaf symptoms (chlorotic and necrotic lesions and malformed leaves) are also sometimes observed in seedlings raised from the fungus-inoculated seeds of Acer saccharum (Picea glauca and Pinus strobes). A new seedling wilt disease of Dalbergia sissoo in India has been identified as Fusarium oxysporum (Harsh et al., 1992). A seedling disease causing collar rot in Azadirachta indica due to Fusarium semitectum has been reported causing 3 to 66 per cent seedlings mortality in 2 to 3 months old seedlings (Uniyal, 1999). Fusarium solani has been found to cause root rot and seedling blight of Azadirachta indica, Eucalyptus camaldulensis and Paraserianthus falcataria ( Sankaran et al.,1986). Acacia koa seedling wilt due to Fusarium oxysporum f. sp. koae has been reported in Gardner (1980).

    Germination Reduction
    The extremely common and numerous mold fungi, are the species of Mucor, Rhizopus, Trichothecium, Botrytis, Penicillium and others, which colonize the surface of several tree seeds and get into the surface tissues. Saprophytic fungi, such as, Aspergillus spp., Mucor spp., Rhizopus sp., Trichoderma sp., and Cladosporium sp. grow on seed coat under favorable conditions, and may invade tissues of the germinating seeds and kill the seedlings. The seed coat microflora could thus, be directly responsible for the weakening of seed vigor, predisposing it to the attack of soil-borne pathogenic fungi. Certain Fusarium spp. isolates and tested, through inoculation on Pinus patula seeds, reduce germination, and cause damping-off of seedlings but do not affect seed germination or seedling growth. Leucaena seeds infected by Colletotrichum graminicola fail to germinate, and if the infection is carried to the nursery, seedlings under moisture stress succumbed to the damping-off disease. Botryodiplodia theobromae, which causes black dry rot in mahagony (Swietania macrophylla) could cause upto to 92 per cent seed deterioration. The causal agent of pitch canker disease of pines, Fusarium subglutinans f.sp. pini has been a new introduction to California (Storer et al., 1998). The pathogen is seed-borne in Monterey pine (Pinus radiata). Seeds from infected branches show up to 83 per cent of infection. Seedling emergence is significantly reduced in infested pine seeds compared with 67 per cent for uninfested seeds. The fungus is more frequent on seedlings originating from diseased branches than from symptomless branches. However, seeds from symptom less branches produced asymptomatic but infected seedlings in Monterey pine.

    Decay and Loss of Viability During Storage
    Storability of seeds is dependent upon temperature, time, relative humidity and method of storage as well as the moisture content of, and initial fungal inoculums on, seeds to be stored. Improper storage of cones cause heating of cones as a result of biological activity and these cones suffer more damage by fungi. The storage containers also affect the seed mycoflora. Storage of achenes of Platanus occidentalis at low temperature (2°C) show no loss in germinability even after 7 months. While at 20°C and 30°C germinability decreases and most fungi on achenes increase with the increase in temperature, relative humidity and time of storage.

    The seeds in Pseudotsuga menziesii cones are safely stored for 225 days under operational conditions and are often free of diseases but when stored after extraction, 56 per cent of them became diseased. Therefore, subfreezing temperatures has been suggested for storing tree seeds at to maintain their germinability of the extracted seeds of Pseudotsuga menziesii, immediately after their removal from the cone to inhibit further fungal activity within the seed.

    Seed Health Testing for Forest Seed

    Most of the forest pathology is found to be confined to seed-borne fungi only. Seed-borne fungi testing include isolation and study of fungi during processing, storage, germination and seedling growth. The seed-health testing methods include direct observation, washing test and incubation methods (blotter and agar plate) for seeds, and seedling symptom test and growing-on test for seedlings (International Seed Testing Association, 1993).

    Some special methods like dilution plate method, ultrasound technique, isozyme patterns, seed tissue excision, seed sectioning, radiography, and ELISA technique have also been discussed. The ISTA recommendations for germination testing, which are more clearly available for agricultural crop seeds, are usually followed as standard practices. Seed size and, sometimes, unavailability of tree seeds in large quantities makes it difficult to use large numbers of seeds in testing. Therefore, it is important to find out (standardize) how many seeds of a tree species should be tested and in how many replicates. For many years, it has been difficult to secure accurate, maximum germination of all viable seeds or achieve the true planting value of forest tree seeds. For example, the dormant nature of the seeds of Abies balsamea, A. fraseri, and other Abies species, which require a moist pre chill treatment of 21 to 28 days or more at 3 to 5°C, together with fungal contamination and growth during the 2-month overall test duration, have been responsible for sometimes erroneous, erratic, or negative germination results. Another problem generally encountered in seed testing involves seed pretreatment. Some scientist use 0.5 per cent sodium hypochlorite solution for 2 to 3 minutes for surface sterilization of diseased red pine seedlings. For testing pathogencity through artificial inoculation of seeds with some seed-borne fungi, different methods for seed inoculations are employed. Several workers have attempted rolling of seeds on fresh fungal cultures, whereas others used spore suspensions. At several platforms, a controversy existed over the method of inoculation and the testing environment, which need to be standardized for different types of seeds. The reduction of germination under conditions of artificial infection does not correspond exactly to the reduction of germination that occurs under conditions of natural infection. Under natural conditions, the various microorganisms on seeds interact within themselves and with the microorganisms present in the soil or growing media. Such an interaction is often even antagonistic, which affects the ability of individual microorganisms to develop rapidly and to infect seeds. Often, in artificial inoculation studies, the conditions for facilitating microbial growth are provided. This suggests a need for testing the pathogenicity of various fungi in natural soils or growing media under greenhouse or field conditions.

    Molecular Techniques for Seed Health Testing
    A number of molecular techniques have been developed or are underway for detection of seed borne pathogen in agricultural crops. Nevertheless, there is need to standardize the molecular and serological techniques, which are more sensitive and quicker for detection of seed borne pathogens in forest seed crops. These techniques may support to screening the material for diseases of significance even in vegetative propagating planting material for sanitary and phytosanitary reasons. Molecular-based identification and phylogeny of Armillaria species has been carried out by Keca et al. (2006).

    Management of Seedborne Infection in Trees

    Seed Collection, Processing and Storage
    Collection of seeds from healthy, disease-free areas or orchards; collection from healthy trees, healthy cones or acorns at the appropriate time; collection from tree and not from ground or from squirrel caches, etc.; transport of cones or seeds in well aerated, clean, dry containers or bags; avoiding damage to seeds during extraction and processing; and use of optimum seed extraction and storage conditions, all need be studied for different forest tree species and considered important for prevention of fungal infections on seeds.

    Surface Treatment
    Although adverse effects on seed germination have sometimes been reported, seed treatment with sterilants to reduce or eliminate fungal contamination has been considered necessary for production of healthy seedlings at several nurseries. For sterilizing conifer seeds with minimal stimulation or retardation to them, an immersion of the seeds in a commercial detergent followed by treatment with 30 per cent hydrogen peroxide has been recommended. Hydrogen peroxide treatment (30 per cent for 45 minutes) improved the total germination in tree seed. In Tectona grandis, surface sterilization helps to control most spermoplane microflora. Sulphuric acid seed treatment reduces seed mycoflora and increases the per cent germination of the seed. Emerging seedlings in such treatments are healthy and are free from storage fungi (Mohanan et al., 2005). In general, hot water and acid treatment enhance seed germination and reduce the number of spermoplane mycoflora and their intensity in tree seeds.

    Chemical Management
    Coating seeds with a repellent against birds and small rodents and a fungicide against damping-off has been a common practice in forest tree nurseries at several places. Although a lot of literature on chemical seed treatment and control of seed-fungi has accumulated, most studies confine to conifers. The sulphuric acid treatment to Araucaria excelsa seeds, which has been prescribed by quarantine regulations against Cryptospora longispora, was found effective in eradicating the seed-borne fungi but the acid was detrimental to seed germination. PCNB dust (70 to 75 per cent) applied to the seed of Balsam, Fraser, and Grand firs gives excellent control of Rhizoctonia solani without any injury to the germinating seedlings. In different tree species (Cedrus deodara, Eucalyptus citriodora, E. hybrid, Pinus roxburghii, P. wallichiana, and Shorea robusta), seed treatment with Brassicol, Bavistin SD and Dithane M-45 as seed dressers could be used to effectively for the control of the most common seed-borne fungi. Effective control of Aspergillus niger on the seeds of Shorea robusta has been found with Bavistin SD or Brassicol seed treatment. The combined use of Benlate (0.15 per cent) and Dithane M 45 (0.15 per cent) has been found to effectively control for several fungi such as, Botryodiplodia theobromae, Colletotrichum gloeosporoides, Fusarium spp., Macrophomina phaseolina, Pestalotia sp., Phoma sp. and Phomopsis sp., on the seeds of Acacia auriculiformis, Albizzia spp., Gmellia arborea, Leucaena leucocephala and several Pinus spp.

    Since detrimental effects of chemical seed treatment on seed germination and seedling quality have also been reported, it is desirable that lower concentrations, which should not be phytotoxic, be tried. Captan at concentrations up to 2,500 ppm did not affect seed germination of Pinus resinosa; however, concentrations of 500 ppm or higher injured roots, stems and cotyledons in young seedlings. Root injury consisted of collapse of root hair cells epidermal cells, and cortical cells; and the cotyledon injury included the collapse of epidermal and mesophyll cells. Ziram and Bavistin have been found effective for management of collar rot in Azadirachta indica ((Uniyal, 1999). Collar rot of Bauhinia purpurea caused by Rhizoctonia solani was recorded for the first time in India. The fungus also attacked cotyedons and caused rotting to varying extent. The disease was effectively controlled by seed treatment with Thiride @ 6g/kg and soil drenching with Dithane M-45 (0.3 per cent) @ 20 ml per tube in root trainers prior to sowing of seeds. The seedling wilt of Dalbergia sissoo, which is severe during summer, and high temperature and moisture favour the disease development is effectively reduced with soil drench with 0.2 per cent Bavistin or seed dressing with 0.2 per cent Topsin-M (Harsh et al., 1992). To safeguard the seeds against spermoplane microflora and also to check the seedling diseases in nursery, Chacko and Mohanan (2002), suggested the use of thiride or captan 2g/kg seeds for D. sissoides seed treatment. The use of Dithane M-45, Bavistin and Captan, a general seed dressing recommendation, for management of seed mycoflora and to improve seed germination and emergence of healthy seedlings in Populus deltoids has been made by Day and Debata (2000).

    Legislation

    Vigorous implementation of the seed laws, such as the Seed Acts and Seed Certification Programmes for quality evaluation and management for forest seed need to be standardized. Moreover, seed collection, seed sampling, extraction, storage and movement need to be standardized keeping in mind the phytosanitary issues.

    Conclusion

    There is an increasing awareness worldwide that unless we intensify efforts at gene conservation, reforestation and intensive forest management, serious depletion of the world’s forests will result. Although reforestation is recognized as an essential activity, an adequate supply of seeds of high quality and high genetic potential is often a limiting factor in many countries. This emphasizes the need for organized seed production and seed research to resolve many problems related to reforestation. Several fungi have been studied on tree seeds; they vary in different host species, in different regions and in different years. Even the detrimental effects to seeds during germination and storage, and to seedlings in nurseries, vary in different host species and environments. With the favorable environment in the tropics, viz. high atmospheric temperatures coupled with high humidity, damage to seeds and seedlings is greater. Biotic factors like squirrel and seedbug damages, and abiotic factors like time and method of collection, shipment, extraction, processing, testing, and storage of seeds, all affect the occurrence of fungi in seeds. Improvement in these practices, use of surface sterilants and/or fungicides and following legislated practices like quarantine will help in the worldwide management of seeds.

    References

  • Ali, M.M.I. and Sharma, J. K. 1996.Impact of seed mycoflora on seed germination and seedling vigour of some important indigenous tree species of Kerela. In: IUFRO Symposium on Impact of Disease and Insect Pests in Tropical Forests, Peechi, 23-26 November 1993. Proceedings. Peechi, KFRI. pp 33-51.

  • Centre for Agriculture and Biosciences International. 2000. Global forestry compendium (CD-ROM). Wallinfford, CABI.

  • Chacko and Mohanan 2002. Development of technology for collection, processing and testing seeds of important tree species of Kerala. Final Technical Report (ICFRE). Kerala Forest Research Institute, Peechi, Kerala.

  • Dey, A. and Debata, D.K. 2000. Studies of leaf spot diseases of Populus deltoids Marsh caused by Alternaria raphani Indian Forester, 126: 1013-1014.

  • Food and Agriculture Organisation of the United Nations. 2000. Corporate document repository, state of world’s forest, Part I: The situation and recent developments in the forestry sector. Rome, FAO. 37p.

  • Gardner, D.E. 1980. Acacia koa seedling wilt caused by Fusarium oxysporum f. sp. koae, f.sp. nov. Phytopathology, 70: 594-597.

  • Harsh, N.S.K.; Tiwari, C.K. and Nath, V. 1992. Fusarium wilt of Dalbergia sissoo Roxb. seedlings. Indian Journal of Forestry, 15(1): 64-67.  

  • International Seed Testing Association. 1993. International rules for seed testing. Seed Science and Technology, 21: 1-259.

  • Keca, N.W.; Bodles J.A.; Woodward, S.; Karadžic, D. and Bojovic, S. 2006. Molecular-based identification and phylogeny of Armillaria species from Serbia and Montenegro. Forest Pathology, 1: 36.

  • Mehrotra, A. 1999. Cotyledon and collar rot of Bauhinia purpurea and its management. Indian Journal of Forestry, 22(2): 177-178.

  • Mehrotra, M.D. 1998 .Rhizoctonia aerial blight - a destructive nursery disease and its management.Indian Forester, 124 (8) 637-645.

  • Mishra, B.M. 2001. Cercospora dehraduni , a new species from India. The Indian Forester, 127: 717-719.

  • Mittal, R.K.; Anderson, R.L. and Mathur, S.B. 1990. Microorganisms associated with tree seeds. Forest Pathology, 30: 305-320.

  • Mohanan, C.; Chacko, K.C.; Chandran, A. and Varma, G. 2005. Seed health problems in tropical forest tree seeds and their impact on seedling production. In: Finnish Forest Research Institute. Working papers. The author. pp.83-93.

  • Pei, M.H. and McCracken, A.R. Eds. 2005. Rust diseases of willow and poplar. Wallingford, CABI. 288 p.

  • Rees, A.A. and Phillips, D.H. 1986. Detection, presence and control of seed borne pest and diseases of trees with special reference to seeds of tropical and subtropical pines. (DFSC Technical Note No. 28). Danida Forest Seed Centre.

  • Salerno, M.I. and Lori, G.A. 2008. Association of seed-borne Fusarium species on Pinus ponderosa with germination and seedling viability in Argentina. Forest Pathology, 37: 263-271.

  • Sanakaran, K.V. and Sharma, J.K.1996 .Diseases of Paraserianthes falcataria in Kerela and their possible control measures. In: IUFRO. Symposium on Impact of Diseases and Insect Pests in Tropical Forests, Peechi, 23-26 November 1993. Proceedings. Peechi, KFRI. pp 142-143.

  • Sharma, J.K.; Mohanan, C. and Maria Florence, E.J.1985. Disease survey in nurseries and plantation of forest tree species grown in Kerela. Peechi, KFRI. 288p.

  • Storer A.J.; Gordon, T.R. and Clark, S.L. 1998. Association of the pitch canker fungus, Fusarium subglutinans f.sp. pini, with monterey pine seeds and seedlings in California. Plant Pathology, 47(5): 649-656.

  • Uniyal, Kamla. 1999. Collar rot in Azadirachta indica and its control. Indian Forester, 125(5): 513-516.

  • Vujanovic, V.; St-Arnaud, M. and Neumann, P.J. 1999. Susceptibility of cones and seeds to fungal infection in a pine (Pinus spp.) collection. European Journal of Forest Pathology, 30: 305-320.


  • Viewpoint: Implementation of Forest Tree Seed Certification and Legislation in India- What Are We Waiting For?

    R.C. Thapliyal*


    Introduction

    I n several countries around the world, where forestry contributes significantly to the overall social and economic well being of citizens, there is growing emphasis on the use of improved seeds in plantations which not only promise higher rate of survival but also enhanced productivity. Keeping this objective in mind, tree seed certification schemes have been initiated for the production, distribution and use of improved seed and other propagation material and legislations enacted in some countries to ensure that only truthfully labeled seed is made available to tree planters.

    The term ‘certified’ implies genetical improvement and aims to make available to working foresters’ seeds, plants and other propagating material of genetic identity and superior quality (Matthews, 1964). Rudolph et al. (1974) in his definition of seed certification emphasized on both the ‘guarantee of seed character and quality, by an officially recognized organization usually evidenced by a certificate, which includes such information as ‘certification category, genuineness of species and variety’ as well as ‘the year of collection, origin, purity, soundness and germination capacity.’ Certification of seed thus essentially involves two aspects viz., the production of seed possessing all qualitative and quantitative traits as desired by the prospective planter and the physical and physiological characteristics of seed which provide a measure of the planting value of a seed-lot.

    Forest Tree Seed Certification in India – a Brief History

    The subject of seed certification in Indian forestry has a long history and has been reviewed earlier (Thapliyal, 2007). Champion (1930) and Dent (1948) were pioneers in emphasizing the importance of ‘seed origin’ and quality of parent trees in plantation forestry in India. Taking a clue from them, an all India symposium at the Forest Research Institute, Dehradun in 1958 recommended for creation of a seed testing and certification agency at the institute. Later, British forest geneticist, Dr. J.D. Mathews developed a tree improvement programme for a few select Indian species which involved development of seed certification procedures, notably for teak, a register of tree and seed sources, seed production areas, etc. was recommended to be maintained. It was proposed that all teak seed used in India be collected from seed production areas SPAs and passed through the seed certification procedure (Kedarnath and Mathews, 1962).

    The subject of forest tree seed certification was revived after a 15 year hiatus in 1976 when the Indo-Danish Project on Seed Procurement and Tree Improvement (IDPSPTI) was taken up. The project took a scheme for the control of forest reproductive material moving in international trade proposed for universal adoption by the ‘Organization of Economic Cooperation and Development (OECD, 1974). The project brought out a publication titled ‘A seed zoning system in India’ (Gopal and Pattanath, 1981) and organized working group meetings with the state forest departments with action plans for execution at state and national level. Provenance trials at national level were taken up for some selected species and lists of selected sources of some priority indigenous species were compiled. Information regarding plus trees, seed stands and seed orchards were also compiled and maintained in standard forms. The project came to an end in 1986.

    In 1988, the National Wasteland Development Board, Ministry of Environment and Forests, Government of India prepared a scheme entitled ‘Central Sector Scheme for Production, Certification, Storage and Distribution of Forest Seeds’, under which funds were provided to interested states for the establishment of ‘seed collection cells’.

    Tree improvement activities were taken up once again in 1993 under the World Bank aided ‘Forestry Research, Education and Extension Project’ (FREEP, 1993-2000). This project, however, was named as the ‘Planting Stock Improvement Project’. Main activities included creation of seed production areas, selection of plus trees, etc. but the major emphasis was on the creation of vegetative multiplication gardens from plus trees. It was in this project that for the first time rouging of undesirable trees was taken up on a large scale in the selected stands for conversion into SPAs.

    Similar developments can be traced in respect of forest tree seed technology. The importance of seed technology in plantation programmes in India was discussed in the Sixth Silvicultural Conference at FRI, Dehradun in 1945 and later by Dent (1948). In the All- India Symposium organized by the Forest Research Institute at Dehradun in 1958 resolution regarding the establishment of a seed testing laboratory at FRI was passed and subsequently first seed testing laboratory in forestry was established at FRI, Dehradun in the year 1962. This laboratory brought out a set of recommendations for the testing of tree seeds of more than 100 species from the Indian sub-continent (Thapliyal et al., 2003).

    Following inputs provided by a paper ‘Organization of Seed Testing Laboratories’ (Thapliyal and Rawat, 1986) presented at the ‘National Seminar on Forest Tree Seed, Hyderabad, 28-30 January 1986 (1986) led to the drafting of a ‘Central Sector Scheme for Production, Certification, Storage and Distribution of Forest Seeds’, as mentioned above. It was through this scheme that funds were made available for setting up seed testing laboratories, apart from seed collection cells, in the state forest departments. Seed testing laboratories were also established at agricultural universities and some institutes through the Research Grant Fund of the Forestry Research, Education and Extension Project of ICFRE (Thapliyal, 1997). A number of training programmes in tree seed technology beginning 1990 were run for the state forest department officials in order to create technical manpower to take up seed collection, processing and routine seed testing in these laboratories.

    Thus over the years, since the visit of J.D. Mathews, the task of marking of plus trees, establishment of seed production areas and seed orchards has been going on unabated (Emmanuel et al., 1992; Gurumurthi, 1992). Two reports emanating from the Indian Council for Forestry Research and Education, (ICFRE), Dehradun listed plus trees, seed production areas, clonal and seedling seed orchards and provenance trials of a number of species undertaken by a chain of ICFRE institutes in various parts and states of the country (Sharma et al., 2002; Katwal et al., 2003). This is in addition to the selection work done by the state forest departments. Seed testing laboratories and seed collection cells in the ICFRE institutes and in some of the state forest departments have also been duly established.

    Present Status of Tree Seed Certification

    According to Barner (1975), a comprehensive forest tree seed certification scheme should encompass the following elements:

    Planning
    1. Preparation of maps showing distribution of important species.
    2. Delimitation of the regions of provenances of these species.
    3. Delimitation of major regions of afforestation and reforestation.
    4. Estimation of demand and supply of seed and plants.

    The Indo-Danish Project on Seed Procurement and Tree Improvement proposed the concept of ‘Seed Zones’ in place of the ’Regions of Provenances’. The data with regard to points 3 and 4 is expected to be available with the state forest departments.

    Implementation
    1. Organization and management.
    2. Classification and approval of sources.
    3. Recommendation for choice of provenances and transfer of reproductive material.
    4. Production and control procedures.
    5. Data recording and documentation.
    6. Marketing of reproductive material.

    Organization and Management
    The following responsible authorities may be appointed for the implementation of a seed certification scheme

    a. Designated authority
    This may be called as certification agency or management committee at central and states level, appointed and be responsible to the government.

    b. Advisory committee
    Constituted by equal representation of foresters, industry, research and seed traders.

    c. A working group on approval of sources
    Appointed by the designated authority, the members of the group include people having practical experience in the fields of silviculture, provenance research, forest tree breeding and genetics, forest mensuration and utilization.

    d. Inspectors
    Should be professional foresters or personnel with special training, acting on behalf of the designated authority.

    The Indo-Danish Project with representation from the Government of India and state forest departments constituted a designated authority, an advisory committee and working groups during the initial project period. On points 3, 4, 5 and 6, scant details are available except may be for teak in some states.

    From the foregoing, it is apparent that some spade work for embarking on a forest tree seed certification programme in the country is in place and the existing gaps and remaining tasks can be accomplished through a seed certification agency whenever constituted.

    Seed Legislation
    Seed legislation has been enacted in most countries around the world to provide guideline and controls to ensure that the purchaser of seed receives a produce that is true to description and of known quality.

    In agriculture, with a view to regulate the quality of seed of any kind or variety to be sold, Seed Act, 1966 was passed by the Parliament which came into force from 1969. Under the Act, the term ‘seed’ includes seeds of crops including edible oilseeds, seeds of fruits and vegetables, cotton seeds, fodder seeds, seedlings, tubers, bulbs, rhizomes, roots, cuttings of all types, grass and other vegetatively propagated material of food crops and cattle fodder. The law provides for initiating comprehensive seed certification of improved varieties and the enforcement of seed control programmes of all seeds sold through the commercial channels for sowing purposes. Under Section 3 of the Act, the central government has constituted a Central Seed Committee to advise the central and state governments on matters arising from the administration of this Act and to carry out other functions assigned to it by or under the Act. This committee is assisted by the various sub-committees at the central level and state sub-committees at the state level in the discharge of its functions. The Seed Act, 1966 was further amended in the year 1972 to provide for the establishment of a Central Seed Certification Board to advise the central government and the state governments on all matters relating to seed certification. Similar organizational set up may be required for the administration of an Act related to forest tree seeds.

    The purpose of seed legislation is to provide legal support to seed certification and is not in itself a part of certification. Legislation places responsibility on the seller to ensure that the seed conforms to the statements made about it or meets certain established standards. For example, if a seed vendor claims to sell ‘certified seed’ provisions under the seed act can be invoked to ascertain whether this seed actually belongs to the said category and really comes from trees in a seed orchard, or from superior (plus) trees in natural stands with artificial pollination. The same will apply to selected seed (seed produced from a SPA) whether the SPA/seed orchard in question maintains minimum standards regarding rouging and an effective pollen dilution zone among other requirements, as laid down under the OECD Scheme

    Apart from the fine details as cited above, the accurate and adequate records about a seed-lot not only enhance the value of seed service but also help seed legislation. All phases of seed operations, from field harvesting to nursery sowing, must be linked by a comprehensive system of records, which on request can be made available for inspection. The most important information about a seed lot which every user would expect to receive with the seed includes the seed lot identity number (Thapliyal, 1994), information on seed origin, year of ripening and an estimate of the quality of seed (viable seed per kg or germinative capacity). Pertaining to germination and purity, there are two main types of seed legislations used, described as ‘minimum standards’ and ‘analytical labeling’. Under the ‘minimum standards’ legislation used in most countries the government sets arbitrary standards for germination and purity, and seed falling below these standards is prohibited from sale. In effect the purchaser is protected from against receiving very poor quality seed. The ‘analytical labeling’ provides specific information on seed quality and the same is printed on seed parcels. This legislation has been enacted in all states of Australia and applies to seed sold in the state for the purpose of sowing (Blackstock, 1987).

    Under the Seed Act, seed testing must be carried out in laboratories duly notified under the Act for labeling requirements. These laboratories should be adequately equipped and staffed with trained personnel to carry out accurate and reliable tests. The seed should also be collected by collectors registered by the seed certification agency and extracted, cleaned and packaged in a registered seed extraction plant and stored in a registered seed storage facility for the satisfaction of the seed purchaser and also for meeting the legal requirements.

    Seed legislation will go a long way in promoting the use of high quality seed in forestry plantations in India but unless the requirements of a comprehensive tree seed certification programme are addressed adequately, the legislation in forestry seed will not serve the desired objectives

    References

  • Barner, H. 1975. The storage of tree seeds. In: FAO/DANIDA Training Course on Forest Seed Collection and Handling, Chiang Mai, 17 February - 13 March 1975. Report. Rome, FAO.

  • Blackstock, J. 1987. Legal aspects of the use of native plant seed. In: Langkamp, P.J. Ed. Germination of Australian native plant seed. Melbourne, Inkata Press.

  • Champion, H.G. 1930. Second interim report on the progress of investigation into the origin of twisted fiber in Pinus longifolia Roxb. Indian Forester, 56 (12): 511-520.

  • Dent, T.V. 1948. Seed storage with particular reference to the storage of seed of Indian forest plants. Indian Forest Records (New Series) Silviculture, 7: 1-134.

  • Emmanuel, C.J.S.K.; Kapoor, M.L. and Sharma, V.K. 1992. Three decades of forest genetics and tree improvement. Indian Forester, 118: 489-500.

  • Gopal, M. and Pattanath, P.G. 1981. Certification of forest reproductive material in India, Revised Scheme, 1979 - Seed Zoning System Followed in India. Hyderabad, Indo-Danish Project on Seed Procurement and Tree Improvement. 25p.

  • Gurumurthi, K. 1992. Baseline studies and basic approaches to tree improvement networking. In: National Network Inception Workshop on Improved Productivity of Man-Made Forests Through Application of Technological Advances in Tree Breeding and Propagation. Coimbatore, 24-25 September 1992. Proceedings. The author.

  • Katwal, R.P.S.; Srivastava, R.K.; Sudhir Kumar and Jeeva, V. 2003. State of forest genetic resources conservation and management in India (Working Paper FGR/65E). In: Food and Agriculture Organisation of the United Nations. Forest genetic resources working papers. Rome, FAO.Kedharnath, S. and Matthews, J.D. 1962. Improvement of teak by selection and breeding. Indian Forester, 88: 277-284.

  • Matthews, J.D. 1961. A programme of forest genetics and forest tree breeding. (FAO/ETAF Report No. 1349). Rome, FAO.

  • National Seminar on Forest Tree Seed, Hyderabad, 28-30 January 1986. 1986. Proceedings, edited by S.N. Rai. The author. 242 p.

  • Organization for Economic Cooperation and Development. 1974. OECD scheme for the control of forest reproductive material moving in international trade. Paris, OECD. Directorate for Agriculture and Food.

  • Rudolf, P.O.; Dorman, K.W. and Hitt, R.G. 1974. Production of genetically improved seed. In: Schopmeyer, C.S. Ed. Seeds of woody plants in the United Sates, Washington, DC, U.S. Department of Agriculture. pp. 53-74.

  • Sharma, M.K.; Singhal, R.M.; Sudhir Kumar and Jeeva, V. 2002. Regional update for India forest genetics resources. In: Twelfth Session of the FAO Panel of Experts on Forest Gene Resources, Rome, 21-23 November 2001. Working papers. Rome, FAO.

  • Thapliyal, R.C. 1994. Identification of forest reproductive material for certification. In: Joshi, N.K. Ed., New trends. Indian forestry. Dehradun, ICFRE. pp. 89-101.

  • Thapliyal, R.C. 2007. Forest tree seed testing and certification in India. In: Seminar on Concepts in Forestry Research, Srinagar, 1-3 November 2006. Concepts in forestry research: Proceedings, edited by N.P. Todaria, B.P. Chamola and D.S. Chauhan Dehradun, International Book Distributors.

  • Thapliyal, R.C. and Rawat, M.S. 1986. Organization of seed testing laboratories. In: National Seminar on Forest Tree Seed, Hyderabad, 28-30 January 1986. Proceedings, edited by S.N. Rai. The author. pp.205-216.

  • Thapliyal, R.C.; Thapliyal, M.; Rawat, M.S.; Nawa Bahar; Naithani, K.C. and Bist, J.P.S. 2003. A handbook for testing of Indian tree seeds. Dehradun, F.R.I. 40p.


    QUALITY SEED PRODUCTION THROUGH HIGH DENSITY SEED ORCHARDS

    Santan Barthwal and Ashok Kumar
    Forest Research Institute, Dehradun 248006


    Introduction

    P rimary objective of an improvement programme is to develop superior planting stock. In commercial deployment of such superior planting stocks, the seed plays a crucial role. Majority of forestation schemes depends on quality seeds and nursery. In forestry, the seed orchards are realistic source of quality seed production. The concept of high density seed orchard is presented to cope up with the growing scarcity of land resources and increasing demand of quality seeds for raising forestry plantations.

    Seed Orchards

    Seed orchard is an area where seeds are mass-produced to obtain the greatest genetic gain as quickly and inexpensively as possible. Seed orchards are the plantations of selected clones or progenies, which are isolated and managed to produce frequent, abundant and easily harvested crops of seeds. Most clonal seed orchards have been established based on some assumptions including (a) all the clones and ramets in the orchard would flower during the same period, (b) have the same cycle of periodic heavy flower production, (c) be completely inter-fertile with all its neighbors and yield identical numbers of viable seed per tree, (d) have the same levels of self-incompatibility and (e) have a similar rate of growth and crown shape as all the other trees. It is almost certain that these assumptions are not virtually satisfied fully. The assumptions of synchronization of flowering and random mating should definitely come true for a seed orchard to be successful.

    There are two types of seed orchards namely clonal or vegetative seed orchard and seedling seed orchard. Clonal seed orchards are those established through the use of vegetative propagules such as grafts, cuttings, tissue culture plantlets, etc. These orchards are the most common type used operationally. The seedling seed orchards on the other hand, are established by planting seedlings followed by rouging the poorest trees, generally leaving only the superior most trees of the best families for seed production.

    There is no rule of thumb as to the number of clones or families that should be raised in the seed orchard. It has been an accepted fact that about 20 genotypes are sufficient to generate a progeny base broad enough to ensure adaptation of orchard strain to normal planting sites, and to avoid serious inbreeding among the genotype within the orchards. This is premised on the condition that the genotypes as much as possible will all flower in equal abundance at the same time, and the localizing of ramets or progenies in the orchard is minimized to avoid inbreeding. For practical reason, it is advantageous to establish more than 20 genotypes to allow for the loss or rejection of stock-scion incompatibility. Strategically, it is wise to include higher number of genotypes in the first generation orchard if progeny test can be conducted to evaluate the genotypes of a younger age.

    Seed orchards are commonly categorized by generation i.e. first, second or more advance generation depending upon how many cycles of improvement a seed orchard has undergone. When seedling seed orchards are the main production populations in a breeding programme, careful planning needs to be done so that advantage can be taken of the fast generation turn over. A basic requirement will be having sufficiently large number of families in the first generation seed orchard to allow for selection and culling in subsequent generations without danger of inbreeding. It must ensure that a large diversity is maintained in each generation by having a representation of at least two thirds of the families of the previous generation (Cameron et al., 1989).

    Recurrent selection for general combining ability is the most commonly used breeding strategy when advance generation breeding is undertaken through seed orchards. A single plantation is established per generation which serves first as a progeny test then as a basis for selection and breeding for the next generation and finally as a SSO. The ultimate aim is to obtain maximum gain per unit of time. Thus it is essential that the improved material obtained through recurrent selection be brought into production orchards as soon as possible.

    First Generation Orchard
    The first-generation orchards are usually established from selections of natural stands or unimproved plantations, most often using individual selection methods. In these orchards, the pedigrees of the parent trees are usually not known. In the first generation orchards the inferior trees after progeny testing are removed by rouging allowing inter mating only amongst the superior trees. The removal of trees for spacing or health reason is simply thinning and cannot be called genetic rouging, because the first generation orchard is started with parents whose genetic worth is not known. The orchards are usually established at a close spacing to allow rouging of poor genotypes, which leave a fully productive seed orchard. Often 50 per cent of the initially established clones are removed. A strategy for developing and testing suitable material for future generations to be designed very early during the first generation breeding programme itself so that broad genetic base is retained and the risk of inbreeding in the advanced generation is reduced. If wider genetic base of economically important traits, such as straightness, wood qualities and growth is to be maintained, it is vitally important to increase genetic variation within the breeding population. The breeding population from which selections are to be made need to be enriched by other material derived from controlled crossings and carefully planned breeding. Each breeding population would effectively provide a base on which successively better production orchards can be established.

    1.5 Generation Orchard
    It consists of taking the best general combiners from a number of orchards similar in geographic backgrounds and bringing them together into a new, improved first generation orchard. These orchards results in excellent genetic gain and produce improved seed because it is composed of the genetically tested genotypes.

    Advance Generation Orchard
    The techniques used in the advance generation orchards is similar to those used in first generation orchards but improved selection methods and designs are used to get greatest gain per unit time. Hence, progeny testing is an essential component of an advanced generation orchard as progenies of the best trees through selection create the base material for next generation breeding. An advanced generation orchard is established from selections made from an orchard where pedigree information and performance of the parents in the test environment is known, unlike an early stage orchard where selections from natural stands are tested. In advance generation orchards for maximum efficiency within family selections and out crossing among the final seed producers, it is necessary estimation of genetic parameters and ranking of families. As inferior individuals are removed in the selection for the next generation, there is less tree-to-tree variation for the important traits in the advancing generations and genetic advance is more dependent on family selections (Barnes, 1995). Selection within family can be reduced as the progenies tend to have greater uniformity and hence the number of individuals per family per se need not be very large. Suitable mating designs, which provide reasonable assessment of the breeding value of parents and produce sufficient genetically distinct families to be used for developing an adequate base for advanced generation orchards.

    Hybridization Orchards
    The occurrence of putative inter-specific hybrids has generated considerable interest in breeding for hybrid production in tree species. Inter-specific hybrids have been reported in pines, eucalypts, acacias and other species when they were introduced to related species. Such hybrids have developed into land races like ‘Mysore gum’ with distinct morphological features and adaptations. Hybrids show best vigour and plasticity when they are introduced to a different environment as new genetic entity with the adaptability of both parent species. Interspecific hybridization is a means of producing a strong heterozygote, which show increased plasticity (Martin, 1989). Crossing should however be performed between species which have an optimum genetic distance and compatibility. Crosses between too close or too distinct individuals will result in inbreeding depression or appearance of abnormal progenies. Apart from tree vigour and adaptability, there are other traits like tree form, resistance to diseases and pests and wood characteristics, which generally have high heritability and can be exploited through hybridization. One of the advantages of creating hybrids is the potential to combine interesting groups of characteristics not found in a pure species (Martin, 1989). Generally F2 hybrids are quite variable but in some species F2 are vigorous as F1. Such F2 hybrids have been used for establishing operational plantations of pines over large areas in Queensland, Australia (Nikles, 1996).

    Spontaneous hybrids also occur in overlapping regions of the species in natural distribution. Heterosis depends to an extent on the genetic diversity of the populations crossed and may be expressed in ideal environments. The heterosis may also result from two or more characters complimenting each other, which is an additive gene effect. Thus, depending on the extent to which the parents differ and the direction of the crosses, certain specific crosses may result in increased hybrid vigour.

    Genetic trials can be used to compare the gains of the second-generation progeny with those of the first generation as well as progenies of individual plus trees of best natural provenances. Montagu et al. (1998) reported 13-21 per cent improvement in growth of second-generation progenies over that of first generation progenies in Acacia auriculiformis. There were, however, several outstanding trees from natural provenance stands whose progeny out performed the second generation progenies of selected mother trees of SSO. This observation shows that vigour increases with improvement in each generation but it is important to include new selections from natural stands as infusions. Seedling seed orchards established with a poor base of inferior local selections have the drawback that the potential gains from out crossing and intensity of selection within the orchard cannot compensate the original inclusion of poor genotypes in the orchard (Montagu et al., 1998).

    In seed orchards establishment, securing full stocking should be the primary concern. Where the field grafting is adopted for establishment, it is advisable to plant two or three seedling stocks in close proximity at each place where successful ramets is desired. This is to allow for grafting failures, culling of incompatible ramets, or selection of best ramets. Orchards established by using potted grafts, balled grafts or seedlings, should adopt a closer spacing to allow for establishment losses.

    Individuals selected for clonal propagation in the orchard should be sexually mature to avoid long delays and great height growth before the orchard trees flower {this is unavoidable in seedling seed orchards and orchards established with rooted cuttings from juvenile sources). Besides providing wider spacing and fertilizing in order to enhance early and abundant flowering, it is imperative for one to learn which environmental factors influence seed production. Thus, some preliminary studies on the suitability of the proposed orchard site for seed production are required.

    Complete isolation of the orchards from inferior pollens of the same or hybridizing species may not be possible. For tropical pines, it is recommended the minimum isolation distance to be 200m upwind from sources of contamination. Effective pollen dispersion distances of hardwoods are not yet well documented. Therefore, the hardwood seed orchards should as much as possible is distantly separated from sources of pollen contamination. Isolation zones are also required to protect orchards from different physiographic regions and between advanced and early generation orchards. However, some species in the same genus need to be isolated from each other because they flower at different times and they will not cross. This is common in pines.

    The size of orchards to be established depends upon the extent of area to be reforested each year and anticipated annual seed yield. It is quite common for initial estimates of the required orchard area to be conservative because of under estimation of future reforestation programme and/or over estimation of seed yield. However, once suitable orchard sites have been developed, it is desirable to reserve large areas in the vicinity to develop later generation orchard, seed extraction plants, nurseries, etc.

    Factors Influence Seed Production

    Physiological plant factors along with environmental condition influence flowering and seed production. Some trees are alternate bearers while some produce seeds every year. The phenology should be well studied for the species of interest for establishing a seed orchard. Some times brief stress may promote flowering in many plant species.

    For some species the reproduction can be hastened by using grafts or cuttings, as the graft retains the physiological maturity needed for flowering. The position of the tree from where the scions are taken also influences the flowering (Schmidt, 1993). The reduction in height of tree is desirable for training of the tree for manipulation of canopy. The grafting of superior scion for tree seed orchards on dwarfing root stock would be required.

    The general requirement in the orchard design is to minimize inbreeding and maximize out crossing (Schmidt, 1993). About spacing of seedlings or clones in seed orchards, it is recommended to have trees 10 m apart, and that accounts for about 100 trees per hectare (Schmidt, 1993).

    High Density Seed Orchards

    Due to land constraints and high population pressure, it might not be possible to have wide layouts for seed orchards and high density seed orchards (HDSO) would replace the conventional orchards. The orchard with high density of trees with intensive fertilizer and water management along with strategic canopy management through tree training will increase quality seed production per unit area substantially. The concept is not very new for horticulture crops where tree canopy is modified to increase the light interception to increase fruit yield. The branches can be trained in a way to increase flowering as well as to practice mechanized collection of fruits and seeds. The intensive management of water and nutrient can be achieved by monitoring of water and nutrient status of the orchards using automated system and their optimum maintenance through drip irrigation. If needed the photoperiodic requirement can also be optimized using artificial lights.

    Pruning is generally recommended in the dormant season and it is advisable to remove branches in such a way to allow penetration of more light into the tree canopy. Higher the light interception percentage better will be the fruiting. For achieving this in fruit crop like Avocado the slender pyramidal canopy is maintained through pruning in New Zealand (Thorp et al.). The method described for Avocado was adapted / modified from that described in apple by Tustin (2000).

    To have high plant density, orchard should have grafted plants of superior genotype on dwarfing root stocks. The trees should be trained from beginning i.e. at the time of planting so as to avoid crown competition and optimizing the light availability and branch density. Controlled crossing can also be achieved easily because of shorter height of the orchards. High density seed orchard offers flexibility in regulating the contribution of individual genotype in seed production. The equal contribution of pollen can be controlled through selective pruning. The desired clones may be allowed to retain larger crown than the less desired ones. Managing genetic diversity of the seed lot can, thus, be controlled through management of flowering in the orchard. Such types of hi-tech seed orchards needs to be managed by highly qualified and well trained mangers.

    Requirements of HDSO
    The high density seed orchard needs two major requirements, first trees producing seeds and second is the technical requirement that includes infrastructure and silvicultural management operations. Trees producing genetically superior seeds are the main component of the seed orchard and rest all revolve around it.

    Ideotype tree for HDSO

  • Should have desired commercial traits needed for its genetic improvement like higher photosynthetic assimilation rate, lower photorespiration, higher water use efficiency, higher nutrient use efficiency, higher biomass, stem straightness, wood and fiber properties, disease / insect pest tolerance and tolerance to environmental stresses.
  • The tree should have profuse fruit and seed producing habit.
  • The tree should respond well to propagation by grafting. This would be required to control the height of the tree in orchard for high density planting and mechanization of seed collection.

  • Technical requirements

  • Commercially available microprocessor controlled irrigation and fertilization system.
  • Sensor based real time soil water and nutrient monitoring system linked to automated irrigation and fertilization system. The optimization of water and nutrient management in orchards may allow significant improvement in yield and environmental protection.
  • Automated pest management system. There are some attempts to develop systems for management of pest in plantation. Silvia-forest management system is a computer application was designed for such purposes (Vallejo et al., 2006). Exclusive software for seed orchards may be developed, which can utilize remote sensing application for monitoring and management of orchards.

  • The utility of high-tech systems for management of orchards have been demonstrated. A spatially variable micro-sprinkler system for orchards was developed and installed for 50 trees in a nectarine orchard with individually addressable micro-sprinkler nodes for each tree. Continuous pressure monitoring allowed fault detection and accurate computation of water discharge by individual micro-sprinklers. The system can precisely control irrigation and fertilization for each and every tree of the orchard (Coates et al., 2005). Monitoring of plantation through satellite has been reported for pine plantation (Joshnston et al., 1997). With advancements in remote sensing technology now it is possible to monitor even one square meter from space.

    For shaping the orchard canopy for better light interception, it is essential to have data on growth and development of canopy of species of interest. Folie et al. (2003) conducted a study to work out modeling of crown growth for tropical forest species. It was reported that trees of 60 cm bole diameter would each require 0.009 ha of growing space with a density of about 107 stems ha-1. Stand basal area converge around 31 m2 ha-1. Such a study would help in planning the layout for the planting in high density seed orchards.

    Such seed orchards can be planned only with tested superior genotypes and clone as gradual thinning of inferior trees is not part of silvicultural operations for HDSO. Replacement of dead and diseased trees should be done as and when required. It is recommended to go for open pyramid training of trees so as to maximize light interception. The other systems of training may be adapted based on the site location of the HDSO and direction of the sun light.

    Applications of Growth Regulators
    There are number of growth regulators that are utilized in horticulture for regulating various stages of plant growth. Auxins are known to promote shoot elongation, cytokinins stimulate cell division, gibberellins promote cell division and elongation. Ethylene a gaseous plant hormone promotes fruit ripening and senescence. Inhibitors of gibberelic acid biosynthesis like phosphone D, amo-1618, cycocyl, ancymidol and paclobutrazol are used to inhibit stem elongation (Salisbury and Ross, 1992). Chemicals like ancymidol, butralin, C8—C10 fatty alcohols, chlormequat chloride are commercially used as plant growth retardants (Fishel, 2006). Gibberellic acids have been used for promotion of flowering in seed orchards. The use of growth retardant (paclobutrazol and flurprimidol) was found to prevent the phototropic curvature of new shoot growth toward increased light intensities in etiolated zinnia seedlings and in silver maple. The growth retardants paclobutrazol and flurprimidol application after pruning reduce the shoot growth and reduced the frequency of pruning operations (Sperry and Chaney, 1999). Such treatments can be utilized for shaping up the canopies in high density seed orchards. Studies on effect of growth retardants on flowering and fruiting would help in deciding the type, concentration, timing and frequency of plant growth regulators in seed orchards.

    Use of Mulching
    Mulching refers to covering of soil surface around plant or tree in a plantation using fabric, plastic metallic, wooden sheets or other material to control weeds and promote tree growth. The mulch prevents loss of moisture from the soil surface, increases the soil temperature (especially when black colour mulch is used), prevents weed growth. Mulches are more popular treatments in fruit than in forestry crops. Weed management through mulching have been reported in many fruit crops (Dale, 2000; Forsella et al., 2003; Starast et al., 2002; Weber, 2003). In orchards of Japanese quince (Chaenomeles japonica), synthetic mulches were found best to control weeds, improve plant growth and yield, and prevented the fruit from getting contaminated by soil (Kviklys et al., 2004). In intensively managed seed orchards, suitable mulch may improve the growth of plants, control weeds and may also promote early ripening of fruits.

    Use of Molecular Markers
    Molecular markers can be utilized for identification of genetically diverse clone for higher genetic gains, estimating genetic contribution and mating system operating in the seed orchard. Ample published literature is available on different kinds of molecular markers, and their applications (Karp et al., 1997; Farooq and Azam, 2002; Spooner et al., 2005; Semagn et al., 2006).

    Conclusion

    Seed orchards that are well-planned and managed, whose clones and families are properly evaluated produce good quality seeds, but their cost in absolute terms is high. Normally the cost involved in developing the required seed orchard is high and is imperative that they be associated with a large reforestation programme. Cost of high density seed orchard with satellite and microprocessor based hi-tech management system (Fig.1) would be still higher, however, considering scarcity of land and water resources and huge requirement of quality seeds it is justifiable to evolve our seed orchard to intensively managed high density seed orchards.

    Fig. 1. Hypothetical design of high density seed orchard with microprocessor based intensive management system

    References

  • Barnes, R.D. 1995. The breeding seedling seed orchard in the multiple population breeding strategy. Silvae Genetica, 44: 81-88.

  • Brown, A.H.D.1990. Plant population genetics, breeding and genetic resources. Massachusetts, Sinauer Associates. pp.145-162.

  • Cameron, J.N.; Cotterill, P.P. and Whiteman, P.H. 1989. Key elements of a breeding plan and temperate eucalypts in Australia. In: Conference on Breeding Tropical Trees: Population Structure and Genetic Improvement Strategies in Clonal and Seedling Forestry, Pattaya, November 28 –December 3, 1988. Proceedings. Oxford, Oxford Forestry Institute. pp. 159-168.

  • Coates, R.W.; Delwiche, M.J. and Brown, P.H. 2005. Precision irrigation and fertilization in orchards. [Available at: http://asae.frymulti.com/abstract.asp?aid=19815&t=2]

  • Dale, A. 2000. Black plastic mulch and between row cultivation increase black currant yields. Hort Technology, 10(2): 307-308.

  • Fishel, F.M. 2006. Plant growth regulator. Florida Cooperative Extension Service, Institute of Food and Agricultural Sciences, University of Florida.

  • Foli, E.G.; Alder, D.; Miller, H.G. and Swaine, M.D. 2003. Modeling growing space requirements for some tropical forest tree species. Forest Ecology and Management, 173: 79-88. 

  • Forcella, F.; Poppe, S.R.; Hansen, N.C.; Head, W.A.; Hoover, E.; Propsom, F. and McKensie, J. 2003. Biological mulches for managing weeds in transplanted strawberry (Fragaria x ananassa). Weed Techology, 17(4): 782-787.

  • Johnston, J.J.;  Weigel, D.R. and Randolph, J.C. 1997. Satellite remote sensing: An inexpensive tool for pine plantation management. Journal of Forestry, 95 (6):16-20.

  • Karp, A.; Kresovich, S.; Bhat, K.V.; Ayad, W.G. and Hodgkin, T. 1997. Molecular tools in plant genetic resources conservation: A guide to the technologies. Rome, International Plant Genetic Resources Institute.

  • Kviklys, D.; Rumpunen, K. and Ruisa, S. 2004. Mulching systems and weed control in Japanese quince (Chaenomeles japonica Ldl.) plantations orchard management in sustainable fruit production. Journal of Fruit and Ornamental Plant Research, 12: 125-132.

  • Martin, B. 1989. The benefits of hybridization. How do we breed for them? In: Griffen, A.R.; Matheson, A.C. Eds. Breeding tropical trees: Population structure and genetic improvement strategy in clonal and seedling forestry. Oxford. Oxford Forestry Institute. pp 79-92.

  • Montagu, K.D.; Nguyen, H.N.; Woo, K.C. and Le Dinh Kha. 1998. The growth of Acacia auriculiformis provenances and seed orchard progeny in Vietnam and Australia. In: International Workshop, Hanoi, 27 - 30 October 1997. Recent developments in Acacia planting: Proceedings edited by J.W. Turnbull, H.R. Crompton and K. Pinyopusarerk. Canberra, ACIAR. pp. 317-321.

  • Nikles, D.G. 1996. The first 50 years of the evolution of forest tree improvement in Queensland. In: QFRI-IUFRO Conference, Caloundra, 27th October- 1st November 1996. Tree improvement for sustainable tropical forestry edited by M.J. Dieters; A.C. Matheson; D.G. Nikles; C.E. Harwood and S.M. Walker. V.2. The author. pp. 50-64.

  • Salisbury, F.B. and Ross, C. 1992. Plant physiology. 4th ed. Belmont, Wadsworth Publishing Company. 682 p.

  • Schmidt, L. 1993. Seed orchards: Guidelines on establishment and management practices. Laguna, UNDP/FAO.

  • Sperry, C.E. and Chaney, W.R. 1999. Tree growth regulator effect on phototropism: Its implication for utility forestry. Journal of Arboriculture, 25: 43-47.

  • Spooner, D.; Treuren, R. van and Vicente, M.C. de. 2005. Molecular markers for gene bank management. Rome, International Plant Genetic Resources Institute.

  • Starast, M.; Karp, K. and Paal, T. 2002. The effect of using different mulches and growth substrates on half-high-bush blueberry (Vaccinium corymbosum x V. angustifolium) cultivars ‘Northblue’ and ‘Northcountry’. Acta Horticulturae, 574: 281-286.

  • Thorp, G.; Woolf, A.; Boyd, L.; Ferguson, I.; White, A. and Everett, K. Avocado canopy management Sustainable production of top quality fruit. [Available at: http://www.avocadosource.com papers/Research_Articles/bundaberg_paper.pdf]

  • Tustin, D.S. 2000. The evolution of central leader apple tree management in New Zealand. Compact Fruit Tree, 23 (3): 83-92.

  • Vallejo, A.; Kanninen, M.; Montero, M. and Viquez, E. 2006. Silvia: A flexible system for integrated plantation management. In: Tewari, V.P.; Srivastava, R.L. Eds. Multipurpose trees in the tropics: Management and improvement strategies. Jodhpur, Scientific Publishers (India). pp. 422-427.

  • Weber, C.A. 2003. Biodegradable mulch films for weed suppression in the establishment year of matted row strawberries. Hort Technology, 13(4): 665-668.


    QUALITY SEED PRODUCTION OF GRASSES AND LEGUMES FOR FOREST GRAZING RESERVES AND DERIVED GRAZING RESOURCES IN INDIA

    N.P. Melkania
    VI/1918, G.B. Pant University of Agriculture and Technology Campus, Pantnagar - 263 145


    Introduction

    I ndia is characterized today by 2.47 per cent (328.76 million ha) of the global geographical area harbouring, respectively, 16.1 and 15.1 per cent human and livestock population with just 1.72 per cent of the world’s forests. Livestock population is the integral component of mixed farming systems practiced in the entire length and bredth of the country. Globally, India possesses the largest livestock population (520.6 million) and accounts world widely for largest number of cattle (16.1 per cent share) and buffalo (57.9 per cent share), second largest number of goats (16.7 per cent share) and third highest number of sheep (5.7 per cent share; Food and Agriculture Organisation of United Nations). Livestock production is predominantly a low input –output system in India. It is primarily practised by marginal farmers, small and medium land holders under moisture deficit/stress and nutrient deficient conditions. As a result of poor availability of productive nutritions fodder and well –managed grazing lands (only 3.4 per cent of the geographical area is under permanent pastures and grazing lands), domestic livestock either subsist on poor quality grasses, legumes and forbs available in degraded grazing lands, forest grazing reserves and non-pasture (waste) lands or are stall fed chiefly on crop – by -residues. Nearly, 60.6 per cent livestock graze in forest reserves (Forest Survey of India, 1996). It is generally agreed that around 30 per cent fodder requirement of livestock at national level is met from forest areas, the removal from the forest, therefore, is to the extent of 145 Mt dry and 178 Mt green fodder annually (Forest Survey of India, 1996). Region – wise, grazing and range forage cultivation is major issue in arid and semi–arid tropics, mountainous region, particularly Himalaya and ecodevelopment projects in protected areas in diverse eco-climates of the country.

    Grazing lands (commonly known as grasslands and rangelands) in India are derived bio-resources from natural forests due to human interventions as deforestation and abandoned cultivation, except the high altitude grasslands in Himalaya, Nilgiris and Palni hills. The bovine pressure is so acute that today almost all types of grazing areas and forest grazing reserves experience overstocking. Although, India ranks first in milk production at global level, the balance sheet of fodder depicts deficiency in the order of 22 per cent for dry fodder, 62 per cent for green fodder and 64 per cent for concentrates (India. Planning Commission, 2002), which is predicted to be 65 per cent for green fodder and 25 per cent for dry fodder in the year 2025 at the current level of growth of forage resources (Indian Grassland and Fodder Research Institute). The shortages would further be aggravated by the adverse impacts of climate change on agriculture production (Sirohi and Michaelowa, 2007) and resultant increase in drought as experienced in 1987 and 1999-2000 (India. Ministry of Statistics and Programme Implementation. Central Statistical Organisation, 2000) in Rajasthan particularly. The emerging dominance of food crops production and growing emphasis on intensive arable cultivation, commercial contract farming and bio-fuel crops cultivation, etc., further reduce land area for forage cultivation on productive abandoned lands and agronomic lands. Therefore, potential scope for livestock husbandry and forage production exists on derived grazing areas and forest grazing reserves.

    Availability of quality seed of range grass and legume species (RGLS) is the single most critical factor to meet the ever-increasing and quality fodder demand of domestic livestock. The demand of quantity seed of RGLS has increased further due to biodiversity/environment – cum – rural livelihood security programmes like watershed development, ecodevelopment, joint forest management, wasteland development, etc. Therefore, all aspects of seed production of RGLS, such as, availability of improved cultivars with potential seed yield, production packages, collection and post harvest processing, storage, marketing and distribution need be looked upon in a holistic manner. This synthesis presents an account of quality seed production of RGLS and limitations experienced under Indian conditions.

    RGLS Quality Seed Production: Institutional Initiatives

    Quality seed acts as a catalyst for realizing potential of all other inputs, such as, appropriate agronomy, fertilization, water, agrochemicals, etc., in farming. Timely availability of quality seed of improved cultivars to the users/farmers at an affordable price is crucial for enhancing and sustaining quality fodder production and profitable economic returns from livestock husbandry. Trials conducted in national agricultural research system (NARS) in India revealed that more than 20 per cent average increase in productivity in various crops can be achieved with the use of quality seed of improved cultivars (Rai, 1998). Research network on forage crops, including RGLS and farm forages, at national level, comprises of Indian Grassland and Fodder Research Institute (IGFRI), Jhansi with its three Regional Research Stations (Avikanagar in Rajasthan for arid region, Darwad in Karnataka for southern plateau and (temporarily at) Palampur in Himachal Pradesh for hill region of Jammu and Kashmir and Himachal Pradesh), All India Coordinated Project for Research on Forage Crops headquartered at IGFRI, Jhansi (with its 18 Coordinated centres in state agricultural universities and non-government organization in (13 agro-climatic regions of India) and Central Arid Zone Research Institute (CAZRI), Jodhpur with its Regional Research Station at Jaisalmer under NARS of Indian Council of Agricultural Research (ICAR) and seven Regional Stations on Fodder Production and Demonstration and one Fodder Seed Production Farm Under Department of Animal Husbandry and Dairying, Govt. of India. Although, organized research on forages in India got impetus by establishment of IGFRI as early as in the year 1962, research on quality seed production got strengthened by establishment of full-fledged Division of Seed Technology at IGFRI under VIII Five Year Plan and through research support from National Seed Project launched by the ICAR in the year 1979. A few of the state agricultural universities, e.g., Tamil Nadu Agricultural University, Coimbatore; CCS Haryana Agricultural University, Hisar, and Punjab Agricultural University, Ludhiana have now established Department of Seed Science and Technology who besides food crops also undertake research on seed production in forages. Unfortunately, no private sector institution presently contributes in research and development of RGLS probably due to focus on commercial agronomic crops, farm forages and horticultural species.

    Quality seed production of forage species was achieved through cooperative network supported by National Dairy Development Board (NDDB) and ICAR under Operation Flood –II and III by establishing nine fodder seed units and Northern Regional Fodder Seed Grid (in the year 1992) with three seed processing units (Singh, 2000). Kerala Livestock Development Board is the only agency in Kerala that organizes production and supply of seed of tropical forage species including RGLS. In West Bengal, forage seed production was organized under ‘Fodder and Livestock Development Programme’ jointly by Animal Resources Development Department and Department of Forests under World Bank sponsored ‘West Bengal Forestry Project’ in ninetees (terminated in December 1997). The forage seed processing centres at Salboni and Jalpaiguri were established. Seed production was organized by involving farmers and forest protection committee(s) for grasses, viz., Setaria sphacelata vars narok, nandi and kajungula; Chloris gayana cv callida and katambra; Cenchrus ciliaris cv cluncurry and molopo; Paspalum notatum; Bothriochloa insculpta cv hatch and B. pertusa; and legumes, viz., Stylosanthes guianensis cv schofield, S. hamata, Desmodium intortum, Macroptilium atropurpureum L 31-85 (D.K. Barat, personal communication).

    Improved Cultivars of Potential RGLS for Quality Forage and Seed Production

    The RGLS grow on extremely degraded lands, prevent soil ecosion, creat favorable micro – environment for introduction of more productive non-graminaceous and leguminous species and provide opportunity of livelihood to illiterate ecosystem people and semi-nomads through cash income by sale of livestock and employment. Although, grass and legume flora are well-represented and distributed in a range of habitats throughout the country [1334 grass species of 261 genera (Karthikeyan, 2005); 1152 leguminous species of 179 genera (richest diversity in Himalaya-488 species), sub-family Papilionaceae contributing 71 per cent to the total leguminous species richness (Rao and Chaudhary, 2002)], only a few of them are recognised as potential palatable nutritious forage species.

    Schima-Dichanthium grasscover occupying the whole semi-arid southern plateau, Satpura and Vindyan area below Jhansi in the south and upto southern Bihar in the east, harbours maximum concentration of leguminous flora, but much of the indigenous legumes are not used as forage due to toxic anti-quality and non-palatability factors. Similarly, because of weedy nature, morphological and biochemical limitations of palatability and poor foliage availability, all grass do not serve as potential forages. The commonly used forage species include Alysicarpus rugosus, Atylosis scraebaoides, clitoria ternacea and Rhynchosia minima among legumes (Singh and Shankar, 2002) and Arundinella nepalensis, Bothrichloa intermelia, B. pertusa, Cenchrus ciliaris, C. setigerus, Chryropogen fulvus, Dichanthium annulatum, Pennisetum flaccidum, P. orientale and Themeda anathera among grasses (Melkania and Tandon, 1988; Melkania and Singh, 1989). The areal extent and green fodder yield of selected potential nutritious RGLS are presented in Table 1.

    Table 1. Areal extent and green fodder yield of selected RGLS in India

    Species Area
    (000 ha)
    Eco Climate Green Fodder
    Yield(q/ha)
    Specific Feature
    Range Grasses
    Cenchurs ciliaris 200 Hot arid and semi-arid 150-400 Potential Species for hot and environ
    C.setigerus 30 Hot arid and semi-arid 100-130 Potential Species for hot and environ
    Chloris gayana - Tropical and Sub-tropical sub-humid and humid 200-350 Potential Species for saline and Alkaline soils and wetlands/marshes
    Chrysopogon fulvus - Tropical and Sub-tropical sub-humid and humid 300-500 Potential Species for rainfed bunds and terrace risers in hills and ravinous lands
    Dichanthium annulatum - Tropical semi-arid 200-350 Potential Species for rainfed bunds and terrace risers in hills and ravinous lands
    Lasiurus sindicus 100 Hot arid 70-250 Potential Species for extremly hot arid regions
    Setaria ancepes 40 Sub-tropical humid 500-900 Potential to grow under Pinus roxburghii
    Range Legumes
    Stylosanthes guiancnsis - Tropical humid and sum-humid 300-550 Compatible to local flora
    S .hamata - Semi-arid 350-450 Potential species for abandoned lands shallow soils and ravinous lands
    S.humilis - tropical sumi-humid 400-500 Compatible to local flora

    The role of improved cultivars of RGLS is important in meeting quality fodder requirement in unit area in unit time besides habitat improvement and carbon sequestration. The improved cultivars for grazing areas and forest grazing reserves need be perennial with profuse tillering, ability to develop tussocks in harsh environ (rainfed, drought prone, frost hardiness) with resilience vigour, leafy, able to survive in poor soil, resistant to insect-pests and ability to compete with local flora- the ruderal weeds in particular. Cultivar development programme in range grasses in India was prominently focussed on potential species of the tropical environment as the semi-arid and arid regions of this climate had been the milkshed and animal husbandry areas for milk-based and meat-based products. Suitable niches for quality seed production and cultivars of RGLS developed and identified potentially suitable for Indian forest grazing reserves and derived grazing resources are described in Tables 2 and 3, respectively.

    Table 2. Suitable niches for quality seed production of RGLS in India (updated following Hazra, 1995)

    Species Niche Location / State
    Range Grasses
    Cenchrus ciliaris Central and south central India Jhansi, Rahuri, Dharwad, Coimbatore, Jodhpur, Jaisalmer, Chadan
    Chrysopogon fulvus Central India and north western Himalayan hill region Jhansi, Rahuri, Pune, Palampur, Solan, Almora, Pithoragarh, Pauri, Chamoli
    Dichanthium annulatum Central and south central India Jhansi, Rahuri, Dharwad, Coimbatore, Jodhpur, Jaisalmer, Chadan
    Festuca arundinacea North western Himalayan hill region Temperate region of Jammu and Kashmir, Himachal Pradesh and Uttarakhand
    Heteropogon contortus Central India Deccan plateau and north western Himalayan hill region Jhansi, Rahuri, Pune, Dharwad, Almora, Pithoragarh, Pauri
    Sehima nervosum Central and south central India Jhansi, Rahuri, Dharwad, Coimbatore, Jodhpur, Jaisalmer, Chadan
    Setaria anceps North western Himalayan hill region Palampur, Almora
    Range Legumes
    Macroptilium atropurpureum Southern plateau Hyderabad, Dharwad and Tiptur
    Stylosanthes hamata, S. glabra Southern plateau Andhra Pradesh, Dharwad, Coimbatore, Rahuri and Pune
    S. hamata cv verano Indo-Gangetic plains of West Bengal Sub-tropical sub-humid climate on new alluvial soil (entisol), e.g., Kalyani
    Trifolium repens North – west Himalayan hill region Temperate region of Jammu and Kashmir, Himachal Pradesh and Uttarakhand

    Technology for Quality Seed Production of RGLS

    Good Agronomic Practices
    In case of range grasses of tropical environ, among the seed hardening chemicals tried at Tamil Nadu Agricultural University, Coimbatore, K2HPO4 1.0 per cent solution was found to have influence on seed germination and seedling growth. Direct seeding of grass species was found the best method for achieving even stand for seed production. In all hard seed coated range legumes, generally, soaking of seeds with concentrated H2SO4 for 4-10 min., followed by washing with running cold water for half an hour before sowing in the field, increases seed germination percentage. Soaking of hard – coated seeds in boiling water alone for 30 – 60 sec. may also increase the early seed germination (Singh and Srivastava, 1990). The agronomic practices recommended for quality seed production of RGLS in India are presented in Table 4. Species-specific details for selected RGLS are described as follows:

    Table 4. Good agronomic practices for quality seed production of potential RGLS in India

    Species Specific feature Sowing time; seed rate (kg/ha); spacing (cm) Fertilization (kg/ha) Seed yield (q/ha)
    Range Grasses
    Cenchrus ciliaris Day neutral, drought resistant, prefers light-medium textured sandy to sandy loam and rainfall 250-900 mm per annum June-July (North), Mar.-Apr. (South); 6-7 for nursery, transplanting after 6 weeks at 60-75 cm distance
    In IGFRI – S – 3108, seed yield is maximum at 30x30 cm spacing in first year and 75x75 cm spacing in subsequent years (Rai, 1981)
    For Marwar anjan and CAZRI-358, seed wet soil ratio 1:4 and sowing 1.0-1.5 cm deep (at 50 cm row spacing) with 0.5 cm soil above seed is ideal. For pelleting, seed rate 2.0-2.5 kg/ha, i.e., 3-5 seed/pellet is preferable (M.P. Rajora, pers. comm.)
    80 N in split doses (40 at sowing and 40 at 30 days after sowing)
    40 N 50 P2O5(50 per cent N at sowing, 25 per cent N at sprouting and 25 per cent N at flowering)
    1.5 – 2.5 Start seed collection 12 – 15 days after flowering
    C. setigerus Prefers sandy light textured soil in 200-700 mm annual rainfall areas June-July (North), Mar.–Apr. (South); 6.7 for nursery, transplanting after 6 weeks at 60-75 cm spacing; pelleting similar to Marwar anjan 80 N in split doses (40 at sowing and 40 at 30 days after sowing), 50 P2O5 1.0-1.25
    Choris gayana Upright stoloniferous drought tolerant grass adaptable to saline soil First week of July; 4-6, Nursery (2-3 kg/ha seed) in first week of June, transplanting of 45 days old seedlings in July immediately after rains at 50 cm row to row spacing 60 N 5.0-6.0
    Chrysopogon fulvus Prefers light soil and rainfall between 500-1500 mm per annum June–July (North), Mar.-Apr. (south); 6-7 in 50x30 cm spacing 90 N (Dwivedi and Kanodia, 1985) 1.0-1.5
    Dichanthium annulatum Decumbent branching, prefers black soil in 600-1500 mm annual rainfall regions June-July (North), Mar.-Apr. (South); 6-8 in row 50-60 cm apart with onset of rains 20 N 0.8-2.5
    Sehima nervosum Prefers red gravel sandy loam to loam soil and 600-1200 mm annual rainfall June-July (North), Mar.-Apr. (South); 6-7 for nursery, transplanting after 5-6 weeks at 60-75 cm distance 80 N 1.0-1.5
    Setaria sphacelata Grows well in heavier soils in high elevational areas in tropics and sub-tropics with 800-1800 mm annual rainfall and loamy to clay loam soil Seed 4.0 kg/ha sown in last week of June at 50 cm row to row spacing and 50 cm plant to plant distance, nursery development (seed requirement 2.0-2.5 kg/ha) preferably in June, transplanting after 6 weeks at 60-70 cm distance 60 N (40 N as basal and 20 N afterwards) 0.5-1.5
    Range Legumes
    Macroptilium atropurpureum Prefers sandy to sandy loam soil in 400-1200 mm annual rainfall areas in tropics and sub-tropics June-July; 2-3 kg/ha seed scarified with concentrated H2SO4 for 25 min., washed thoroughly and thereafter sown at a depth of 15-25 cm in 1.0 row spaced plot 15-20 N and 50-60 P2O5; molybdonated SSP (4-5 q/ha) and KCl (1.22 q/ha) applied at sowing; cutting at 15 cm height maintains vigour 1.5-2.5
    Stylosanthes guianensis Prefers medium to heavy rainfall tropics (900-1600 mm annual rainfall) and loam to silty clay loam soil, tolerant to low fertility 5-7 kg/ha seed inoculated with local strains of Rhizobium to be sown in last week of November at 15 cm depth in 0.5 m row by drilling or broadcasting 15 N and 50 P2O5, SSP 100-150 kg/ha applied at sowing; harvest crop 25-25 cm below the top, sun-dry for 4-5 days and thresh for seed 15-20
    S. hamata Prefers sandy loam to loam soil and rainfall 600-900 mm per annum June-July (North), Mar.-Sept. (South); 7-14 kg/ha seed broadcasted on soil surface in 50 cm row spacing 15-20 N, 20 S and 20-30 super phosphate; higher seed yield with 450 kg/ha SSP on sandy loam red soil (Singh,1981); harvesting preferably in mid July 3.0-4.0
    S. humutis Adapted to light textured soil of low fertility, also grows successfully in cracking clay soils Similar to S. hamata Similar to S. hamata 7.5-10.0
    S. scabra Prefers sandy to sandy loam soil and 300-600 mm annual rainfall June-July (North); Mar.-Sept. (South); 7-10 kg/ha seed broadcasted on soil surface in 50 cm row spacing 15 – 20 N and 50-60 P2O5 3.0-4.5

    Setaria (Setaria sphacelata)
    Grass planted at 50 cm row to row distance gives higher seed yield over that planted at 75 and 25 cm distance. Seed yield equivalent to 30 N/ha is obtained when intercropped with Desmodium tortosum with additional leguminous fodder yield (Singh and Srivastava, 1990). S. sphacelata intercropped with Desmodium tortosum IL 1568 increases seed yield and saves 20-30 kg N/ha with additional yield of leguminous fodder (Indian Grassland and Fodder Research Institute).

    Buffle grass(Cenchrus ciliaris)
    Pre-treatment of seeds with 0.2 per cent KNO3 improves seed germination in C. ciliasis and C. setigerus (Indian Grassland and Fodder Research Institute). Crop raised from 8w old seedlings results in higher seed production than that raised from 1y old seedlings and new tillers from old tussock. Transplanting from first to third week of July under normal monsoon conditions gives higher seed yield over delayed transplantings. Seed yield increases with increasing levels of N upto 80 kg/ha. Foliar sprays of urea alongwith KNO3 @ 2.0 kg/ha each provide high seed yield, compared to 2.0 kg each of MgSO4, ZnSO4 and single superphosphate (Singh and Srivastava, 1990). Application of Azospirillum and Azotobactor significantly increases seed yield by 16 and 11 per cent, respectively, compared to no application of biofertilizers. Foliar spray of panchagavya and ghomutra (10 per cent each), KNO3 (2.0 per cent, 4.0 per cent and 6.0 per cent solution) and GA3 (50 ppm, 100 ppm and 150 ppm solution) at spike initiation stage, increases seed production in C. ciliaris and C. setigerus. Thiourea (0.50 per cent solution) increases significantly (46 per cent) seed yield (1.62 q/ha) in C. ciliaris and spray of panchagavya increases seed yield by 16.1 per cent (5.2 q/ha) in C. setigerus over no treatment. Application of these chemicals, also increases 1000 – seed weight (IGFRI, 2006). Foliar application of NAA alongwith K and DAP at flower initiation stage in C. ciliaris and C. setigerus increases seed setting, seed – retention and seed yield (Indian Grassland and Fodder Research Institute)

    Studies conducted at Tamil Nadu Agricultural University, Coimbatore (V. Krishnasamy, personal Communication) recorded removal of inhibitors by soaking seeds of C. ciliaris in water for 48 h. The dormancy is overcomed if the seeds are treated with GA 250 – 750 ppm, KNO3 0.75 per cent or Thiourca 0.75 per cent. Cultivation of C. ciliaris (var CAZRI 75) under rainfed conditions in Kamataka at 45 cm row spacing and fertilization with 40M40P2O5 kg/ha (20 per cent dose of N and complete dose of P2O5 as basal and 20 per cent dose of N as top dress) yielded 72.3 per cent higher seed than the lower P level (seed yield 27.8 kg/ha). It is recommended that during first year of establishment, the crop can be grown for seed production only, while in subsequent years, depending on rainfall, the grass can be cut at six to 12 weeks after monsoon for green fodder and the ratton can be continued for seed purpose (Purushottam et al., 1995).

    Dharafor goria(Chrysopogn fulvus)
    Spacing 50 x 50 cm provides superior seed yield, compared to wider and narrower spacings. Fertilization studies indicate that the grass responds positively only upto 30 kg N/ha (Tomar, 1998).

    Stylo(Stylosanthes hamata)
    Stylo seed has certain inherent germination related problems, such as hard seededness (30-70 per cent) alongwith high proportion of immature seeds and seed dormancy, resulting in poor crop stand. Drying freshly harvested seed in galvanized iron trays in summer months (April -May) increases seed germination by 1.5 to 2 times.

    Dry seed treatment with Bavistin @ 0.2 per cent is recorded best in controlling anthracnose disease (Colletotrichum gloeosporioides) resulting in better germination and seed yield. Scarification with concentrated H2SO4 @ 200 ml/kg of seeds for four min. has been found promoting seed germination. Soaking seeds in GA 50 ppm or KNO3 0.25 per cent for three hours also found effective in improving germination of seeds (V. Krishnasamy, personal communication). Summer sowing of stylo is better than the monsoon sowing for seed production. In monsoon season, first week of July sown crop gives higher seed yield over delayed sowing. Seed rate @ 10 kg/ha is recorded better than five and 15 kg/ha. Phosphorus application @ 48 kg P2O5 /ha doubles seed yield over no phosphorus application (Singh and Srivastava, 1990; Tomar, 1998).

    Sward of S. hamata cv. verano sown in April –May in Indo-Gangetic plains of West Bengal at 40 to 60 cm row to row spacing with fertilization @ 80 kg P2O5/ha, defoliated in July and left thereafter for seed production, yields 140 q/ha green fodder and 7.0 to 9.0 q/ha seed with additional benefit of nitrogen in soil (A. Karva, personal communication). Studies conducted at Mahetma Phula Krishi Vidyapeeth, Rahuri (Desai, 1985) recorded seed yield potential of S. hamata, two to three q/ha in the first year and 8.0 to 10 q/ha in subsequent years. However, seed production after second year requires reseeding and fertilization to maintain vigour to optimal level. Growing S. hamata for dual purpose (green fodder and seed) reduces seed yield which is usually recorded 2.0 to 3.5 g/ha.

    Studies conducted on S. scabra (Srikaanth, 1994) suggest following technological innovations for quality seed production:

  • October to January (preferably November) is optimum and ideal time for collection and sowing of seeds. Grader using BSS 16 x 6 sieve is ideal for recovering quality seeds.
  • Seeds with creamy coat possess higher germination and vigour potential. Soaking seeds in GA 100 ppm or KNO3 0.25 per cent solution for three hours improves germination percentage.
  • Seed germination is highest for seeds harvested at 21 days after anthesis.
  • Highest recoverable amount of seeds could be shelled out with paddy dehusker with its rotating discs having a clearance adjustment of 0.2 mm and 0.3 mm as optimum to realize higher shelling percentage (52 and 56 per cent, respectively)
  • Scarification with concentrated H2SO4 at 200 ml/kg of seeds for four minutes improves germination by reducing the hardening. Scarified seeds retain better viability when treated with thiram @ 2 g/kg of seeds and 0.2 g/kg of carbonyl and stored in 700 gauge polybags.

    The demand of stylo seed for use in wasteland development and land reclamatiion projects (dates backs to 1974) triggered seed production by resource poor small farmers in Anantapur district, Andhra Pradesh as stylo (S. hamata and S. scabra) compares favourably with other crops in terms of inputs required and economic returns under marginal cropping conditions. The seed yield ranges from 50 to 60 kg/ha in the first year to 80 to 100 kg/ha in subsequent years. As the seed is harvested at crops maturity, the viability and germination are, therefore, usually very good. The major contaminates are weed seeds, small pebbles and dried parts of flower spikes which could be removed by mechanical cleaning before sale (Ramesh and Pathak, Memio).

    Siratro(Macroptillium atropurpureum)
    Seed treatment with 100 ppm GA3, hydration for 18 h and dry dressing of thiram and osmo conditioning in PEG solution (-10 bars) increases germination, field emergence and yield of seed in M. atropurpurcum (and S. hamata and C. ciliaris). Seed yield is also increased by foliar application of 100 ppm NAA and 50 ppm MH at 35 days after sowing and flower imitation stage. Climbing support of Sesbania sesban increases seed yield (61.1 kg/ha) as compared to no support (45 kg/ ha; Indian Grassland and Fodder Research Institute).

    White clover(Trifolium repens)
    It combines well with native grasses in sub-temperate and temperate environ, and contributes a considerable quantity of nitrogen in a sward besides improving yield and quality of forage. Seed set is recorded more in lower or first formed florets (1.82 seed/floret) than the upper or late formed ones (1.47 seeds/floret). The seed set is maximum (average of upper and lower florets 2.11 seeds/floret) under partial shade (sun light available for four hours/d only) than the complete light or complete shade (Sareen, 2000). The 1000 seed weight is also maximum (0.554 g) under partial shade. The ovule sterility, lack of pollination and post fertilization abortion are responsible or low seed set is this range legume.

    Management of Biotic Stress
    Almost all important forage RGLS are attacked by insects, bacteria, fungi, virus and nematodes. Tropical grasses, viz., Dichanthium annulatum and Sehima narrosum, and temperate species of Festuca, Lolium, Phalaris and Poa are susceptible to rust, particularly species of Puccinia and Uromyces. Leaf spot caused by Pyricularia spp. is major destructive disease of tropical grasses, viz., C. ciliaris and C. setigerus (Pandey et al., 1995). Anthracnose caused by Colletotrichum gloeosporioides is the most serious foliar disease of Stylosanthes where seed serves as primary source of infection. Root rot caused by Sclerotium rolfsii is another important disease of this range legume. Package of practices for control of diseases for seed production in most of the RGLS are wanted. Recommended measures for managing diseases in selected RGLS are described in Table 5.

    Table 5. Recommended measures for controlling diseases in selected RGLS

    Species Disease Causal organism Protection practice
    Range Grasses
    Cenchrus ciliaris Smut Sorosporoum cenchri S. penneseti Solar heat or hot water treatment, dry seed treatment with Carboxin 0.25 per cent
    C. setigerus Ergot Sphacelia spp. Use of Zriram or Zrineb + copper oxyehloride 0.25 per cent
    Smut Sphacelotheca panjabensis Seed treatment with fine powdered sulphur or Agrosan GN before sowing @ 5 g /kg and 2 g/kg., respectively
    Dichanthium annulatum Leaf spot Curvularia and Ropogonis Spay of 0.20 per cent Zineb, Maneb and Nabam + ZnSO4 at initial appearance
    Smut Entylome obesum Seed treatment with Agrosan GN beore sowing @ 2 g/kg
    Rust Puccinia duthie Spray of Zineb or Dithane M-45 @ 3 kg/ha
    Ergot Sphacelia spp. Use of Zineb or Ziram and copper oxychloride 0.25 per cent
    Range Legumes
    Macroptilium atropurpureum Mosaric Virus Rouge out the affected plants, use Muvacron or Parathion 250 ml /acre
    Stylosanthes guianensis Damping off Piricularia spp. Seed treatment with Metalaxyl 0.3 g/kg seed and captan
    S. humalis Anthracnose Colletotrichum gloeosporioides Seed treatment with Benlate 6.0 g/kg seed and use of blue copper 2.5 kg/ha


    Limitations Experienced in RGLS Seed Production

    Technological Constraints
    Quality seed production in grasses and legumes is an exacting and specialized task and express unique limitations, compared to food grain crops and tree species. The process of seed production is governed by a multitude of genetic, environmental and physiological factors. Apomictic reproduction system in grasses hinders breeding efforts to develop improved cultivars. Knowledge about how to manipulate them, as seed crops due to prolonged period of growth and flowering, is limited. Further low multiplication ratio and high cost of production make RGLS seed production and processing a difficult and costly process. Range of species which can produce seed efficiently to meet the diverse ecological conditions of the country is limited to Schina-Dichanthium cover (found across the whole peninsular India), e.g., Stylosanthes hamata, C. ciliaris and to a lesser extent S. scabra and C. setigerus, and Dicanthium-Cenchrus – Lasiurus cover (northern parts of Gujarat, Rajasthan and Aravelli ranges).

    Poor seed production in RGLS is generally related to shy seeding tendency and poor translocation of water-soluble sugars from vegetative to reproductive parts (high amount of sugar remains either in crown region in legumes and stem portion in grasses), poor translocation mechanism of carbohydrates, low productive potential of late developed flowers, non-synchrony of flowering, prolonged flowering, flower drop, uneven pod maturity and pod shattering, and loss of plant nutrients due to defoliation/cutting, low priority to RGLS, compared to farm forages (like sorghum, berseem, etc.) in cultivation, harvesting crop before reaching seed stage and long duration and high cost of collection of seeds especially in uneven maturity species are the major management problems that add to low seed production. As the improved varieties are basically bred for higher forage yield, these cultures are genetically low seed yielder.

    Indeterminate Growth
    Most of the RGLS have indeterminate growth habit. The non – synchrony in vegetative and floral growth causes low partitioning of photosynthates development and seed yield.

    Uneven maturity
    On account of uneven maturity in reproductive parts, the pod dehiscence and seed shedding results in low seed production even after several rounds of seed collection in a season.

    Blank seed
    In range grasses particularly, major part of the collected seeds is empty and only nearly 30 per cent seeds remain viable.

    Site effect
    Since quality of seed produced depends on photoperiod, thermoperiod, humidity, soil texture, structure, reaction and moisture, better seed yield could be achieved if the seed crop is grown in the site(s) of species –specific requirements.

    Non-synchronised flowering
    Forage grasses have the tendency of non-synchronised flowering within the population and within a panicle. Seeds would reach maturity in some spikelets while some of the spikelets would be in anthesis. Under such situation, fixing the optimum stage of harvest becomes difficult. Defoliation, adjusting irrigation and fertilizer application are essential to achieve synchronised flowering. Non – synchronised flowering has got an adaptive advantage. If unfavourable weather or stress occur, only a portion of the reproductive unit will be affected and others will help in perpetuation.

    Poor seed setting
    Weather condition during flowering has a significant role to play. Dull, cool and rainy period if coincides with flowering, will result in poor seed set. In forage legumes, less activity of pollinators may cause poor seed set, owing to lack of pollination. Internal plant regulation also sometimes leads to ovular abortion in legumes.

    Seed shedding
    Most of the forage species are perennial and flowering extends over a long period. The flower which opens first will mature earlier than others and the ripened seeds are not retained on the panicle.

    Harvesting
    Shattering causes loss of seeds during harvesting operations. Non-uniform maturity results in incomplete recovery of seeds during threshing. Immature seeds possess high moisture and are susceptible to crushing while overdrived seed sustain cracking injuries during threshing.

    Seed processing
    Uneven maturity again poses difficulty in processing of seeds as the drying will not be uniform. Seeds with diverse maturity exhibit differences in size and so grading is difficult which causes substantial rejection loss. Seeds also contain attached glumes and chaffy materials.

    Seed dormancy
    Dormancy in forage grass seeds is caused by tight glumes and inhibitors present either in glumes or in seeds. Forage legumes invariably possess hard seeds necessitating scarification treatments before sowing.

    Harvest index
    A negative correlation is recorded between fodder and seed yield in forages in general as most of the photosynthates are diverted towards production of vegetative parts and very little is utilized for economic production, i.e., seed. Harvest index in forages, thus, is extremely low (Table 6); grasses have poorest harvest index (> 4 per cent), followed by range legumes (> 12 per cent) and cultivated forages (12 – 35 per cent). Further, emphasizing more on biological yield in varietal development than the economic yield have also resulted further decline of harvest index.

    Table 6. Harvest index of major Indian tropical RGLS

    Range grasses Harvest index Range legumes Harvest index
    Dichanthium annulatum 3.2 Stylosanthes viscosa 50.0
    Chrysopogon fulvus 2.0 S. humilis 8.3
    Heteropogon contortus 2.0 S. scabra 7.3
    Cenchrus setigerus 1.7 S. hamata 5.0
    Sehima nervosum 1.0 S. guienansis 2.6
    Cenchrus ciliaris 0.8 Macroptilium atropurpureum 1.1
    Setaria sphacelata 0.6

    Organizational Constraints
    While the large quality of RGLE seed for programmes like watershed development, ecodevelopment, compensatory revegetation and habitat development for wild animals in protected areas demands large tracts of land or seed production (maintaining isolation distances), most of the seed production institutions in the public sector do not have enough land available and have priority of land for seed production of food crops and/or arable forages. Thus, instead of producing seed by proper planting and monitoring, RGLS seed is collected through hand harvesting or sweeping from existing natural degraded grazing resources by rural poor, women and children in particular. The demand for seed of RGLS is spatially and temporally specific and raised primarily by public sector. There is no centralized authority for demand forecasting, supply of seed and monitoring of quality.

    The development of research on improved cultivars of RGLS for seed production has been slow. Only two institutes under ICAR viz., IGFRI, Jhansi and CAZRI, Jodhpur are involved in breeding programmes on range grasses and legumes. Further, of the cultivars released, L. sindicus (CAZRI – 130 -5) and C. setigerus (CAZRI – 76) got popularity (Baumann and Turton, 1998). Private sector does not contribute on RGLS as these are non – profitable economically, compared to food crops and cultivated forage crops like sorghum, maize and peral millet. A major constraint in the development of new cultivars of RGLS has been lack of feed – back from growers into the research process. Criteria, viz., resilience under drought conditions, easiness in cutting, storage properties and palatability to all kinds of livestock could not receive emphasis like forage yield. Further, except at few locations, most of the trials on RGLS are conducted under farmland conditions which are distinctly different than grazing resources existing under real-life situations. Seed production research on RGLS of high elevational grazing resources of Himalayan region (> 2000 m asl) is yet almost untouched subject.

    Future Perspective

    The concern about degradation of terrestrial ecosystem leading to disappearance of grazing resources and gradual desertification, environmental degradation and investment in wasteland development and watershed management in India has primarily generated an appreciable demand for (quality) seed of grass and legume species adapted to forest and non-forest range and wasteland conditions. Organizations, viz., Department of Agriculture (soil conservation, animal husbandry and watershed development projects), Department of Forests (soil conservation, participatory forestry and ecodevelopment projects), National Afforestation and Ecodevelopment Board, Ministry of Environment and Forests, Ministry of Defense (animal farms and eco-task force), National Wasteland Development Board of Ministry of Rural Development and Non-governmental organizations working in the area of revegetation and rural development are now potential market for quality seed of RGLS. But the research and development efforts in regard to production and availability of quality seed are lagging behind. Shortage in supply of breeder seed alone in forages has been reported as high as 80 per cent (Singh and Khanna, 1995). At times, quality of seed supplied has also been questioned by the indenters. Review of existing seed certification standards in regard to RGLS and development of new standards and standardization of seed testing protocols are still at nascent stage. Creation of nodal authority for organized production and distribution of forage seed at central and state level is wanted. The private sector need be motivated for partnership in research and development for quality seed production of RGLS.

    Social acceptance becomes the foremost criterion in recognition of a cultivar and production technology by the society. As the RGLS are region-specific, research need be based on regional preferences considering the socio-economic factors, indigenous traditions and customs of the regionous people, pastoralists and semi-nomads. The cost effectiveness in terms of finance and maturity duration need be recognised as one of the major criteria while researching technological options for RGLS seed production. The seed yield potential need be considered as a criterion like forage yield and crude protein yield in varietal development programme on RGLS. The future research should be able to meet the following challenges:

  • Development of synchronous and non-shattering type cultivars of RGLS.
  • Genetic improvement with respect to seed set in Trifolium repens.
  • Assessment of genetic purity, marker(s) identification and characterization of RGLS using DNA finger printing and isozyme analysis.
  • Development of cost effective and eco-friendly package of practices for management of production (agronomic practices like seed rate, spacing, application of micronutrients and growth regulators, appropriate time of harvest, etc.) and protection (sanitary measures for controlling pre-and post – harvest losses (due to insects, pathogens and rodents) of quality seed of RGLS.
  • Development of seed collecting devices (considering employment opportunity of the farmers), seed driers and seed separators and post harvest technologies for minimizing mechanical damage and maximizing quality of seed during processing, storage, transport and distribution.

    Acknowledgements

    This synthesis in an attempt to provide a well-informed and intimate account of opportunities for quality seed production of potential grass and legume species of forest grazing reserves and derived grazing resources in India. I am grateful to Drs. S.B. Tripathi, R.K. Bhatt, A.K. Roy, U.S. Mishra (all from IGFRI, Jhansi); M.S. Yadav and M.P. Rajora (CAZRI, Jodhpur); D.C. Kotoch and Naveen Kumar (CSK HPKV, Palampur, Himachal Pradesh); J.P. Yadavendra (GAU, Aanand); J.S. Desale (MPKV, Rahuri); S. Purushottam (UAS, Bangalore); A.K. Fazlullah Khan, A. Gopalan, V. Krishansamy, Manon Mani (all from TNAU, Coimbatore), J.P. Singh and Rajendra Singh (NDDB, Anand), K. Krishnan (KLDB, Thiruvananthapuram); A. Karva (BCKV, Kalyani) and. D.K. Barat (Animal Resource Development Department, Govt. of West Bengal, Kolkata) for valuable discussions on the subject while my working with the ICAR as Project Coordinator (Forage Crops) during 1998 to 2003. Professional support extended by the Director, IGFRI, Jhansi and Director General, Forest Survey of India, Dehradun is thankfully acknowledged.

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    EVALUATION OF SEED QUALITY DURING MATURATION FOR EFFECTIVE COLLECTION

    Maitreyee Kundu
    Tropical Forest Research Institute, Jabalpur- 482 021


    Introduction

    S eed maturity has critical implication for seed quality in terms of their germinability and vigour. Pre-harvest factors such as degree of seed maturity influence germinability and vigor, which in turn, affect the potential longevity of seeds (Justice and Bass, 1978). If the information on seed quality in relation to seed maturity is available, immature seeds can also be harvested to overcome certain contingent situation.

    There are several reasons to develop techniques for collection of immature seeds:

    a. To extend the collection season: The short period between seed maturity and dispersal may be an excessive demand on the availability of seasonal labour and in some areas unfavorable weather conditions in the collection period may aggravate this situation. Lengthening the period available for collection permits better organization of the collection and allows skilled personnel to pick more of the crop. It can be particularly valuable in research, where a large number of seed lots have to be collected from widely scattered localities. Griffin (1990) used the technique in his study of Douglas fir provenances.

    b. To minimize loss of seeds to dispersal/predation: Some fruits are prone to serious predation/dispersal at the early fruit stage. In Africa, many acacias and other legumes suffer from serious attack by bruchid beetles. Bruchid larvae develop within the seeds. Sometimes, more than 50 per cent of the seeds are infected with bruchids when seeds are mature (Doran et al., 1983). Drupes of Maesopsis eminii are removed by hornbills, which may raid a tree in a few days, especially if the fruit crop is small. Flocks of weavers, fruit bats or migratory birds may completely empty a tree for fruits in a matter of hours. Premature collection also applies to many small-seeded, wind-dispersed species with a short maturation season, in which seeds are widely scattered once mature (Gray, 1990). For example, seeds of Ailanthus excelsa are usually picked before maturity since fully mature fruits are liable to lose most of their seeds as soon as attempts are made to collect them (Khullar et al., 1991).

    c. To avoid development of dormancy: Some types of dormancy develop only during the late maturity processes (Mayer and Poljahoff-Mayber, 1982). If seeds are to be sown immediately after collection, pretreatment can sometimes be avoided if the seeds are collected early. An example is physical dormancy in Leguminosae, which develops as a result of dehydration of the seed coat. Seeds are germinable before the seed-coat becomes impermeable and early collected seeds, thus, need no or only a very mild pretreatment.

    d. To reduce pre-processing damage: Fermentation of fleshy fruit pulp may damage the seeds. If depulping is not possible in the field or immediately after collection, fermentation risk may be delayed by collecting fruits early.

    e. To salvage immature seed collected inadvertently: Untrained collectors of seeds often being picking fruits and cones too early in the year before they are fully mature. Artificial ripening provides a method for handling this material.

    Maturity Indices

    The optimal time for harvest is when a large amount of viable, germinable seeds can be collected. This stage of seed can be marked as mature stage. Knowledge of physical changes of seeds and attainment of germinability and seed vigor is essential to determine the best collection time. Generally, fruit/seeds are harvested according to the indicators of maturity (termed as maturity indices) based on physical parameters of the fruits/seeds/cones. Seed collectors have long been aware that most mature and immature fruits and seeds can be distinguished in a number of ways i.e. by color difference, increased firmness or brittleness, decreased moisture content, specific gravity, by changes in physical dimensions or development of abscission zones. Forest tree fruits generally achieve near maximum full size relatively early in the ripening season so that further changes in physical dimensions as full maturity reached is usually too small to be of much use.

    Dehiscent fruits often develop visible aperture structure prior to dispersal i.e. lines where the fruit will eventually split open, e.g. dehiscence lines of the operculum of Eucalyptus capsule.

    For species where harvesting can be done without significant loss of seeds and where the trees can be kept under observation, collection may be postponed until natural dispersal has commenced, hence dispersal in itself is an indication of seed maturity. This can be observed either directly, on opening of dehiscent fruits of falling seeds, or indirectly on frugivores visiting trees with animal dispersal. The majority of seeds of species with a relatively short maturation season are normally mature when the first ones are released. Typically, however, seeds of poor quality (e.g. underdeveloped or infected) are shed before the main crop (Owen, 1995).

    Direct examination of the seeds is applicable for species with relatively large seeds. Cutting tests reveal the development of the embryo and the firmness of seed coat and endosperm. The endosperm, where present, should be firm and not milky. The size of the embryo varies with species.

    Sometimes, biochemical parameters such as crude fat, reducing sugar, minerals, starch, soluble nitrogen and protein nitrogen are used as maturity indices. Bonner (1971) produced a list of chemical constituents for 37 broad leaf species in the Southern United States. But this study on biochemical indicators so far, was unable to clarify the concept of seed maturity.

    Seed Maturation

    During seed development, three broad stages are generally recognized. The first stage is characterized by fertilization, cell division and histodifferentiation of all major tissues (Raghavan, 1986). During the second stage a massive accumulation of storage proteins, lipid and starch occurs, reflected by an increase in the seed dry weight. The maturation stage follows, culminating in the arrest of reserve deposition and desiccation. It is important to determine the time from this physiological developmental stages, when the seed attains maximum quality i.e. the seed attains maximum germination percentage and seedling growth. Controversies regarding this point are related to the enormous diversity of reproductive and adaptive strategies observed in plants (Bewley and Black, 1985). According to Harington (1972), maximum seed quality is achieved towards the end of the filling period, when the seeds reach maximum dry weight. The hypothesis had been checked in various agricultural species, such as Glycine max L. ( Tekrony and Phillips, 1980). However, in other species, maximum seed quality is reached sometime after attainment of maximum dry weight as reported in Vicia faba L. and Lens culinaris Medik (Ellis et al., 1987) and many other species (Demir and Ellis, 1992). Ellis and Pieta (1992) introduced the term ‘mass maturity’ to distinguish the time of maximum dry weight accumulation from that of physiological maturity. Another type of maturation had been recognized in seeds – morphological maturity: morphologically mature seeds have a fully developed embryo and endosperm and other structures. The relative development of the embryo was considered as a good indicator of maturity in some cases (Finnis, 1950; Katsuita, 1975), when the seed attained maximum seed germination and seedling vigour before seed fall. This idea had been contradicted by others (Zasada, 1973), where morphological maturity was achieved before physiological maturity i.e. germinability. The two types or stages are not mutually excluding, since change in physiological ripeness usually accompanies embryo development. It is also important to note from different studies that physiological maturity may not be achieved after the achievement of mass and morphological maturity.

    Rediske and Nicholson (1965) observed that even in late cone collections, when organic accumulation was complete and the embryo had fully elongated, the seeds would not germinate until a period of after –ripening had elapsed. They, thus, described two phases of ripening. First was a period of organic accumulation in the seeds, indicated by a rise in seed weight and an increase in the radiographic density of the megagametophyte tissue; this phase at the end of which the seeds were barely germinable, was referred to as maturation. A second phase, a period of after-ripening after the apparent organic accumulation had ceased, was required to permit germinability to develop. This second phase was achieved by artificially storing intact Abies procera cones for several weeks, before the seeds were extracted.

    Thomas et al. (1973) noted three phases in the maturation of sycamore maple (Acer pseudoplatanus L.). These were described as (i) a period of growth during which fresh and dry weighty increased; (ii) a temporary cessation of weight increase followed by a marked accumulation of organic compound including lipids, carbohydrates, proteins and membrane bound ribosomes, with protein becoming the main storage product and (iii) a period of desiccation. At the end of the second phase, that is before desiccations began, the embryo had no capacity for germination, even though it was morphologically mature. Only during the desiccation phase did the embryo begin to display any germinability and that only in response to chilling.

    Orthodox Seed Maturation
    The seeds that are capable to withstand desiccation and whose longevity is improved with decrease in storage moisture content down to 5 per cent or less are called orthodox (Roberts, 1973). Generally, these type of seeds dry slowly on the plant during maturation (maturation drying) during which, it develops desiccation tolerance. Dormancy is imposed in this stage in some species especially in legumes. The onset of desiccation tolerance after dry mass accumulation marks the earliest time during development of orthodox seed when harvest and rapid artificial drying for achieving physiological maturity is feasible (Long et al., 1981). It has been suggested that seeds for genetic resources conservation might the collected whilst seed moisture contents are quite high (rather than delay collection) in order to reduce pre-harvest deteroration in the quality of the seeds collected and to minimize problems in the timing of collection in harsh and extensive terrains.

    Recalcitrant Seed Maturation
    Recalcitrant seeds are desiccation sensitive both during development and after they are shed. These seeds are shed at high water contents and viability of the seeds rapidly decreases, if moisture content of the seed decreases after a certain limit (lowest safe moisture content). Most of these types of seed accumulate dry mass up to full maturity and there is no or very short maturation drying phase. Desiccation tolerance increases with the progress of maturity and the highest is found in the fully mature seeds (Finch-Savage, 1992; Kundu and Sett, 2005). Since the seeds have short viability and no dormancy, late collection is unsuitable since seeds are easily lost to germination or deterioration (Berjak and Pammenter, 1996). Even the storability of recalcitrant seeds is better in mature seeds in comparison to immature ones (Chien and Yang, 1997; Kundu and Sett, 2005).

    Artificial Ripening
    Huss (1951) reported that some immature Pinus sylvestris seeds in Sweden ripened after extraction during several months of storage in a ventilated cellar. Wakeley (1954) suggested that slightly immature cones, those collected only one to two weeks earlier than recommended, of the major southern pine species (Pinus eliottii, P. taeda L., P. palustris Mill, P. echinata Mill., etc.) may mature after harvest if properly stored. Oliver (1974) reported that White fir cones collected 4 weeks before seed fall can be artificially ripened and these cones yielded seed which germinated as completely and speedily as stratified seed from mature cones.

    From the different studies on seed maturation and artificial ripening, it is clear that there seems to be a point in the development of fruit prior to which the seeds cease development and after ripening procedure will be successful, if they are detached from the parent plant, regardless of how the seeds are handled after harvest. This is because the food reserve necessary for continuing seed development are not available in fruit. Once this critical point has been reached, seed may be after-ripened with proper care. In most of the dehiscent orthodox species, the practical method of assessing maturity is the desiccation tolerance test, as it has long been known that the physiological capacity that marks maturity in this type of seeds is the attainment of the ability to withstand desiccation. Though this technique cannot be applied in field conditions, it would need less laboratory facilities than biochemical method. Once the desiccation tolerance is achieved by the immature seeds, they can be harvested and artificially ripened through drying methods.

    Pre-Curing
    Pre-curing denotes the procedure during which fruits are kept moist for a prolonged period before extraction. Pre-curing has two objectives: a. to promote after-ripening of immature fruits, and b. to ease extraction of seed where rapid desiccation may cause extraction problems, in extreme cases case-hardening. The critical stage after which after-ripening is possible varies among species and must be based on experience. During the natural maturation process, the water supply to the maturing fruit is regulated though the pedicel and to the seed through the funicle. Continuous evaporation prevents overheating of the embryo while it still has high moisture content. Maintenance of high moisture level and avoidance of drastic increase in temperature should simulate these conditions during after-ripening.

    Only fruits that are not mature should be after-ripened. If the seed lot varies in maturity, as can usually be judged from the appearance of the fruits, the fruits should be sorted prior to processing; small under-developed fruits are always discarded as they have not attained the capacity to after- ripen. Fully mature fruits go directly to the next step in the processing chain, e.g. extraction; an intermediate portion may consist of mature size fruits capable of after-ripening during a pre-curing period. For example, in Malaysia pods of Acacia mangium are separated into three classes according to colour viz. greenish brown, brown and black. Greenish brown pods are after ripened for 120 hours, brown pods for 72 hours and black pods go directly to extraction by kiln drying (Bowen and Eusebio, 1982). In Kenya, seeds of Azadirachta indica, Thevetia peruviana and Ximenia americana are after-ripened for 2-3 days after collection (Ahenda, 1991).

    After-ripening typically takes from a few days to a few weeks. The environment during this period is important in order to control the physiological process. The fruits are kept at normal air temperature, in the tropics between 20o and 30oC. The moisture level is initially high but gradually reduces during the process. The high moisture level during pre-curing makes the seed susceptible to fungal attack. Proper ventilation limits that risk, but carries a risk of desiccation.

    After-ripening is concluded when the fruits have reached full maturity. In most dry fruit types, the maturity indices of naturally matured fruits will hold for after-ripened fruits as well. However, since after-ripened fruits are not normally exposed to sunlight, maturity colour of fleshy fruits may be slightly different from naturally ripe ones, and the fruit flesh may remain relatively firm. To judge when maturity has been reached, examination of seed development and the embryo may be carried out by a cutting test.

    Seed extraction of several tropical pines can be greatly facilitated by pre-curing, especially for early collected cones. In an experiment of early collected cones of Pinus elliottii, the number of seeds extracted per cone increased from 0 to 60 for cones pre-cured for 1 and 5 weeks, respectively. When cones were collected one month later, the number of seeds extracted increased from an average of 27 after 1 week’s pre-curing to 82 after 5 weeks pre-curing (McLemore, 1975).

    Pre-curing of relatively dry fruits may be conducted while the fruits are stored in containers of bags. Alternatively, the fruits may be spread in a thin layer (one fruit thick of large fruits) on concrete floors or in trays. In the latter case, it is easier to control the environment during the pre-curing period. Desiccation may be controlled by regularly spraying with water. Spraying is gradually reduced as the fruits reach full maturity.

    Pre-curing of fleshy fruits should always be carried out while the fruits are spread out. That allows moisture from softening fruits to drain off, the fruits should preferably be pre-cured in open trays. If the fruits attain maturity at different rates, it may be necessary to normally remove mature fruits in order to avoid decomposition or fermentation of the pulp while pre-curing is still going on.

    The process of pre-curing is basically the same whether used for after-ripening or to ease extraction. However, since the latter is a mainly physical process, it is less dependent on a physiologically optimal temperature. Consequently, the temperature regime is less critical. For example, for cones it may be advantageous to gradually raise the temperature during the later stages of the process (Morandini, 1962).

    Disadvantages
    There are some potential problems in collection of immature fruits or seeds.

    a. More resources needed: After-ripening needs space, manpower and other resources. Premature collection implies that extraction must be postponed until the seeds are fully mature, which excludes field extraction for reduction of bulk.

    b. More laborious seed collection and extraction: Immature seeds are collected from the trees. As abscission zones have not yet developed, shaking of branches will rarely be sufficient for releasing fruits and the more laborious cutting of fruits or fruit bearing branches will be necessary. Extraction of seeds from artificially ripened fruits is often more difficult than from naturally matured ones. Problems of opening dehiscent fruits and loosening of fruit pulp are frequent, especially in fruits collected very early.

    Conclusion

    As the afforestation program continues to increase the demand of more seeds, it is necessary to expand the collection season. Artificial ripening has been applied to more and more seeds collected earlier in the ripening season. Yet it is difficult to identify the maturation point when the seeds should be collected for after-ripening and also the method of after ripening has not been standardized for many species especially from the tropical origin.

    Even the definition of maturity is not clear from the studies done so far on seed maturity. Some have related maturation to peak germination, others have preferred to relate with morphological and physiological maturity. Morphological maturity i.e. full development of embryo is a key stage for many species, the development of maximum germinability usually occurs after the embryo has become fully elongated. For several species, relative development of the embryo is becoming accepted as an indicator of the time when seeds can be picked and successfully ripened artificially. Physiological maturity or the completion of organic accumulation within the seeds is another key stage in the maturation process, though morphological and physiological maturity occurs concurrently at different rates.

    In some other species, even though organic accumulation has been shown to be complete, the embryo is fully developed, yet the seeds require a so-called after ripening period before full germinability and better seedling growth can be realised. This third stage during which desiccation with some biochemical changes occur in the seed and some of the seeds attain desiccation tolerance in the phase. This stage may be associated with a change in the state of embryo dormancy or degree of delayed germination. For this reason for some species, the problem of breaking dormancy in mature seeds so far exceeds the problem of handling immature seeds, some of which will germinate readily, so much so that proposals have been made to collect and immediately sow large quantities of green seeds.

    The crucial point of collecting immature seeds is to determine the critical point after which seeds can be collected and artificially after ripened without hampering seed quality i.e. maximum viability, absence of abnormal seedlings, rapid germination and better seedling growth. This requires both experience and experiments in which physiological development of the seed is related to one or more physical characters of the fruit or seed.

    Acknowledgments

    The author is grateful to Dr. A.K. Mandal and Mr. Aditya Kumar for inspiration and encouragement.

    References

  • Ahenda, J.O. 1991. Processing of pulpy seeds. In: 1st National Tree Seed Workshop, Nairobi, 1-5 July 1991. Kenya Forestry Research Institute. pp. 114-126.

  • Berjak, P. and Pammenter, N.W. 1996. Recalcitrant (desiccation-sensitive) seeds. In: Olesen, K. Ed. Innovations in tropical tree seed technology. Humleback, Dandia Forest Seed Centre. pp. 14-29.

  • Bewley, J. and Black. M. 1985. Seeds: Physiology of development and germination. New York, Plenum Press.

  • Bonner, F.T. 1971. Chemical contents of southern hardwood fruits and seeds. USDA Forest Service. 3 p.

  • Bowen, M.R. and Eusebio. T.V. 1982. Seed handling practices: Four fast growing hardwoods for humid sub-tropical plantations in the eighties. The Malaysian Forester, 45(4): 534-547.

  • Chien, C. and Yang, Ja-ju. 1997. Effect of seed maturity on storability of Litsea acuminata seeds. Taiwan Journal of Forest Science, 12(3): 369-372.

  • Demir, T. and Ellis. R.H. 1992. Changes in seed quality during seed development and maturation in tomato. Seed Science Research, 2: 81-87.

  • Doran, J.C.; Turnbill, J.W.; Boland, D.J. and Gunn, B.V. 1983. Handbook of seeds of dry zone acacias: A guide for collecting, extraction, cleaning and storing the seed and for treatment to promote germination of dry zone acacias. Rome, FAO.

  • Ellis, R.H.; Hong, T.D. and Roberts, E.H. 1987. Comparison of cumulative germination and rate of germination of dormant and aged barley seed lots of different constant temperatures. Seed Science and Technology, 15: 717-727.

  • Ellis, R.H. and Pieta Filho, C. 1992. The development of seed quality in spring and winter cultivars of barley and wheat. Seed Science Research, 2: 9-15.

  • Finch-Savage, W.E. 1992. Seed development in the recalcitrant species Quercus robur L.: Germinability and desiccation tolerance. Seed Science Research, 2: 17-22.

  • Finnis, J.M. 1950. Seed maturity in doghlas-fir. British Columbia Forest Service. 8p.

  • Gray, R. 1990. Professional seed collection, In: Sowing the Seeds, Direct Seeding and Natural Regeneration Conference. Proceedings. The author. pp. 135-140.

  • Griffin, A.R. 1990. Effects of inbreeding on growth of forest trees and implications for management of seed supplies for plantation programmes. In: Bawa, K.S. and Hadley, M. Eds. Reproductive ecology of tropical forest plants. UNESCO. pp. 355-374.

  • Harrington, J.F. 1972. Seed storage and longevity. In: Kozlowskii, T.T. Ed. Seed biology. V.3. New York, Academic press. pp. 145-245.

  • Huss, E. 1951. Alterations in viability during the storage of cones and seed. Skogen, 38: 151-154.

  • Justice, O.L. and Bass, L.N. 1978. Principles and practices of seed storage. Washington DC., USDA. 289 p

  • Katsuita, M. 1975. Embryo development and germinability of Pinus thumbergii Parl. and P. densiflora Sieb et Zucc. Bulletin of the Tokyo University Forests, 67: 105-134.

  • Khullar, P.; Thaplial, R.C.; Beniwal, B.S.; Vakshashya, R.K. and Sharma, Ashok. 1991. Forest seed. Dehradun, ICFRE. 409p.

  • Kundu, M. and Sett, R. 2005. Develoment of Aquilaria agallocha Roxb. Seeds: Acquisition of germinability, desiccation sensitivity and storage response. Indian Journal of Plant Physiology, 10: 362-366.

  • Long, S.R.; Dale, R.M.K. and Sussex, I.M. 1981. Maturation and germination of Phaseolus vulgaris embryonic axes in culture. Planta, 153: 405-415.

  • Mayer, A.M. and Pljakoff-Mayler, A. 1982. The germination of seeds. New York, Pergemon Press.

  • McLemore, B.F. 1975. Collection date, cone storage period affect southern pine seeds yields, viability. Tree Planters Notes, 26(1): 24-26.

  • Morandini, R. 1962. Forest seed handling, equipment and procedures, I: Seed production, collection and extraction. Unalsylva, 15(4): 1-15.

  • Oliver, William W. 1974. Seed maturity in white fir and red fir. Washington DC., USDA. 12 p.

  • Owen, J.N. 1995. Constraints to seed production: Temperate and tropical forest trees. Tree Physiology, 15: 477-484.

  • Raghavan, V. 1986. Embryogenesis in Angiosperm. A developmental and experimental study. Cambridge, Cambridge University Press.

  • Rediske, J.H. and Nicholson, D.C. 1965. Maturation of noble fir seed: A biochemical study. Weyerhaeuser Forestry Paper, 2: 1-15.

  • Roberts, E.H. 1973. Predicitng the viability of seeds. Seed Science and Technology, 1: 515-527.

  • Tekrony, D.M. and Phillips, A.D. 1980. Effect of field weathering on the viability and vigor of soybean seed. Agronomy Journal, 72: 749-753.

  • Thomas, H.; Webb, D.P. and Wareing, P.F. 1973. Seed dormancy in Acer maturation in relation to dormancy in Acer pseudoplatanus L. Journal of Experimental Botany, 24: 958-967.

  • Wakeley, P.C. 1954. Planting the southern pines. Washington DC., USDA. 233p.

  • Zasada, J.C. 1973. Effect of cone storage method and collection date on Alaska white spruce (Picea glauca) seed quality. In: International Symposium on Seed Processing, Bergen, September, 1973. Papers. IUFRO. pp. 1-10.

    INDIA TODAY’S EVALUATION –

    One of the Finest Institutions of the Country: Forest Research Institute, Dehradun

    The British Empire was founded on wood-first the tall mast timber used in its warships which conquered the world and later, the teak sleepers on railway lines that knitted them.

    The establishment of a dedicated Forest Research Institute (FRI), a little over a century ago, was the culmination of its imperial obsession with wood. Since then, Asia’s Mecca of forestry has become much more than India’s sole university dedicated to forestry. Located on a sylvan 1,200-acre wooded campus in Dehradun, it is a gigantic vault that holds all the data about India’s forest wealth, past and present. The FRI’s xylarium is a library of 20,000 wood specimens of trees from around the world. The 3.3 lakh plant specimens, some over two centuries old, form among Asia’s largest herbariums, and its collection of two lakh forest insects are among the largest in the world. Nowhere in the world are all 13 disciplines of forestry- from genetics and tree propagation to forest pathology –housed under on roof, a splendid Greco-Roman structure. Declared a university 17 years ago, it is a hub of education which churns out nearly 130 scholars as also the forest officers and rangers, from its forest academy, who protect it. “The FRI has contributed to research in forestry and environment, from basic forestry knowledge and technology, to advanced studies in frontline thrust areas like biotechnology, bio-fuels and nano forestry,” says FRI Director S.S. Negi. At a time when the twin threats of global warming and fast depleting fossil fuels occupy maximum bandwidth, the institute is assured of a relevance that runs parallel to the wealth it studies.

    Courtesy: India Today, 30(34), 2008: 70-71


    PATTERN OF SEED DISPERSAL AND REGENERATION IN DIPTEROCARPUS INDICUS: AN ENDEMIC TREE SPECIES OF WESTERN GHATS

    B.N. Sathish*, Syam Viswanath**, C.G. Kushalappa*, and S. Raghavendra*
    *University of Agricultural Sciences, College of Forestry, Ponnampet
    ** Institute of Wood Science and Technology, Bangalore-560 003


    Introduction

    S eed dispersal is widely believed to be a fundamental element of plant population ecology and has received much scientific attention in the last decades (Howe and Smallwood, 1982; Herrera et al., 1994; Fuentes, 2000; Nathan and Muller-Landau, 2000). It is a crucial ecological process that maintains the diversity of plant communities (Christian, 2001; Webb and Peart, 2001), linking spatial patterns of parent plants and their descendants (Harper, 1977; Schupp and Fuentes, 1995; Nathan and Muller-Landau, 2000) Information on rates and spatial patterns of seed dispersal is increasingly recognized as important for understanding the maintenance of species diversity in plant communities (Tilman, 1994; Hurtt and Pacala, 1995; Clark et al., 1999). Most wind-blown seeds fall near the parental tree, but long-distance dispersers are disproportionately important at both evolutionary and ecological scales of time and space (Nathan and Muller-Landau, 2000; Nathan, 2001). Long-distance dispersal helps to maintain metapopulations and gene flow in fragmented and changing habitats like those humans produce habitually (Cain et al., 2000). Dispersal interacts with competition and predation, as well as with other aspects of plant community dynamics (Green, 1983; Horn, 1991; Shigesada et al., 1995). Long-distance dispersal determines rates of invasions, epidemics, range expansions, and responses to climate change (Clark, 1998; Clark, Fastie, Hurtt et al., 1998; Clark et al., 1999; Higgins and Richardson, 1999; Neubert and Caswell, 2000). To examine the consequences of seed dispersal for patterns of plant recruitment, the spatial patterns of dispersed seeds must be quantified and their determinants understood (Nathan and Muller-Landau, 2000). Dipterocarpus indicus Bedd. is a massive evergreen tree which dominates in the lower elevational evergreen forest of western Ghats Very little studies have been conducted with respect to seed dispersal and the distribution patterns of the seedlings. Being one of the globally endangered and endemic tree of the Western Ghats, it is important to understand seed dispersal, seedling distribution pattern, etc., which, in turn decide the population structure of such an important endemic and threatened species for its conservation in the near future.

    Materials and Methods

    The study was conducted in the year 2006 in the tropical low elevational evergreen forests of Makutta (N 12ş 05' 52.3, E 75 ş 42' 35.6, 338 m elevation) in Kodagu district, which is located in central part of Western Ghats. The forests in this area are dominated by tree species like Dipterocarpus indicus, Kingiodendron pinnatum, Vateria indica, Sterculia alata, etc. Ten Fruiting trees of Dipterocarpus indicus, with almost uniform age and height were randomly selected in the study area. Near each tree, sample plots of 1 x 1 meter were laid at every five meters starting from the base of the tree (considering it as 0 m) in four directions. In each plot, all the seeds of Dipterocarpus indicus trees were collected and size in terms of diameter, wing lengths, width and presence of tell tale holes as a result of insect predation/infestation in seed were measured. In addition to it, all the recruits of Dipterocarpus indicus were also recorded in the same plots.

    Results

    It was found that, the seeds have spread over up to 25 meters from the base of the tree. Majority of the seeds were aggregated near to the base of the tree and very few of them were dispersed up to 25 meters as shown in the Fig. 1. The very unique feature of the seeds of this particular species was that most of the seeds were infested by insects evidenced by the presence of holes on the seeds. Majority of the infected seeds were found near the base of the tree, while as we move away from the tree, there were very few infested seeds as shown in the Fig. 2. The number of recruits in the same plots was very low at the base of the tree and increased as a function of distance from the tree (Fig. 3) that means that even though there were very few seeds away from seed source, they were free from insect infestation and majority of the seeds were viable and mainfested as yound recruits (Fig. 4).

    Fig. 1. Variation in seed density at different distances from the seed source

    Fig. 2. Proportion of healthy seeds in total seed count at different distances from the seed source

    Fig. 3. Recruitment pattern of Dipterocarpus indicus at different distances from the seed source

    Fig. 4. Proportion of recruits and healthy seeds in seed density plots at various distance from tree base

    Discussion

    In nature there is vast variation in the mode of seed dispersal it plays a critical role in population and community dynamics of plants (Harper, 1977; Howe and Smallwood, 1982; Schupp and Fuentes, 1995; Nathan and Muller-Landau, 2000). Major sources of variation include internal (parent-controlled) factors, such as seed morphology and height of release and external factors that influence the performance of the dispersal agent(s).

    In general, most seeds are dispersed over short distances; only few travel far from the source (Cain et al., 2000). Similarly in our study most of the seeds of Dipterocarpus indicus seeds were aggregated or clumped near the base of the tree. Although both short- and long-distance dispersal can be generated by stochastic effects, these types of dispersal often result from different mechanisms (Nathan and Muller-Landau, 2000). Moreover, the implications of recruitment from short-distance dispersal could be fundamentally different from those of long-distance dispersal. Short-distance dispersal of the vast majority of the seeds generates a small-scale pattern that has critical implications for local recruitment (Janzen, 1970; Connell, 1971; Schupp and Fuentes, 1995; Nathan and Muller-Landau, 2000). A basic feature of this pattern, is the rapid decline in seed densities with distance from the source (Harper, 1977; Willson, 1993), which leads to strong sibling competition and intensive seed predation in the vicinity of adult trees (Janzen, 1970; Connell, 1971). Because dispersal distances are considerably shorter within dense forests than in open landscapes, Janzen-Connell effects are likely to be more powerful in forests. Even though the total number of seeds near source was very high, most of them were infested by insects. Janzen (1971) argued that the high seed densities within forests could reach a level at which predators become satiated, thereby increasing seed survival and favoring selection for reduced dispersal distances.

    References

  • Cain, M.L.; Milligan, B.G. and Strand, A.E. 2000. Long-distance seed dispersal in plant populations. American Journal of Botany, 87: 1217-1227.

  • Christian, C.E. 2001. Consequences of a biological invasion reveal the importance of mutualism for plant communities. Nature, 413: 635-638.

  • Clark, J.S. 1998. Why trees migrate so fast: confronting theory with dispersal biology and the paleorecord. American Naturalist, 152: 204-224.

  • Clark, J.S.; Fastie, C.; Hurtt, G.; Jackson, S.T.; Johnson, C.; King, G.A.; Lewis, M.; Lynch, J.; Pacala, S.; Prentice, C.; Schupp, E.W.; Webb, T. and Wyckoff, P. 1998. Reid’s paradox of rapid plant migration: Dispersal theory and interpretation of paleoecological records. BioScience, 48: 13-24.

  • Clark, J.S.; Silman, M.; Kern, R.; Macklin, E. and Hille Ris Lambers, J. 1999. Seed dispersal near and far: patterns across temperate and tropical forests. Ecology, 80: 1475-1494.

  • Connell, J.H. 1971. On the role of natural enemies in preventing competitive exclusion in some marine animals and in forest trees. In: den Boer, P.J.; Gradwell, G.R. Eds. Dynamics of populations. Wageningen, Centre for Agricultural Publishing and Documentation. pp. 298-312.

  • Fuentes, M. 2000. Frugivory, seed dispersal and plant community ecology. Trends in Ecology and Evolution, 15: 487-488.

  • Green, D.S. 1983. The efficacy of dispersal in relation to safe site density. Oecologia, 56: 356-358.

  • Harper, J.L. 1977. Population biology of plants. London, Academic Press.

  • Herrera, C.M.; Jordano, P.; López-Soria, L. and Amat, J.A. 1994. Recruitment of a mast-fruiting, bird-dispersed tree: Bridging frugivore activity and seedling establishment. Ecological Monograph, 64: 315-344.

  • Higgins, S.I. and Richardson, D.M. 1999. Predicting plant migration rates in a changing world: The role of long-distance dispersal. American Naturalist, 153: 464-475.

  • Horn, H.S. 1991. Some causes of variety in patterns of forest succession. In: West, D.C. ; Shugart, H. H. and Botkin, D.B. Eds. Forest succession: Concepts and applications. New York, Springer Verlag. pp. 24-35.

  • Howe, H.F.; Smallwood, J. 1982. Ecology of seed dispersal. Annual Review of Ecology and Systematics, 13: 201-228.

  • Hurtt, G.C. and Pacala, S.W. 1995. The consequences of recruitment limitation: reconciling chance, history and competitive differences between plants. Journal of Theoretical Biology, 176: 1-12.

  • Janzen, D.H. 1970. Herbivores and the number of tree species in tropical forests. American Naturalist, 104: 501-528.

  • Janzen, D.H. 1971. Seed predation by animals. Annual Review of Ecology and Systematics, 2: 465-492.

  • Nathan, R. 2001. Dispersal biogeography. In: Levin, S.A. Ed. Encyclopedia of biodiversity. V.2. San Diego, Academic Press. pp. 127-152.

  • Nathan, R. and Muller-Landau, H.C. 2000. Spatial patterns of seed dispersal, their determinants and consequences for recruitment. Trends in Ecology and Evolution, 15: 278-285.

  • Neubert, M.G. and Caswell, H. 2000. Demography and dispersal: Calculation and sensitivity analysis of invasion speed for structured populations. Ecology, 81: 1613-1628.

  • Schupp, E.W. and Fuentes, M. 1995. Spatial patterns of seed dispersal and the unification of plant population ecology. Ecoscience, 2: 267-275.

  • Shigesada, N.; Kawasaki, K. and Takeda, Y. 1995. Modeling stratified diffusion in biological invasions. American Naturalist, 146: 229-251.

  • Tilman, D. 1994. Competition and biodiversity in spatially structured habitats. Ecology, 75: 2-16.

  • Webb, C.O. and Peart, D.R. 2001. High seed dispersal rates in faunally-intact tropical rain forest: Theoretical and conservation implications. Ecological Letters, 4: 491-499.

  • Willson, M.F. 1993. Dispersal mode, seed shadows, and colonization patterns. Vegetatio, 107-108: 261-280.

    World’s Largest Seed

    The world’s largest seed comes from the coco-de-mer palm (Lodoicea maldivica), native to the Seychelles Archipelago in the Indian Ocean. Although it belongs to a different genus from true coconut palms (Cocos), this enormous seed is often called the ‘double coconut’. A single seed may be 30 cm long, nearly 0.9 m (three feet) in circumference and weigh 18 kg. It should be noted here that the largest seed does not have the largest embryo. In fact, palm seeds are mostly composed of endosperm tissue and generally have relatively small embryos.

    Available at: http://waynesword.palomar.edu/ww0601.htm#seed


    SEED GERMINATION STUDIES IN DIFFERENT SEED SOURCES OF AZADIRACHTA INDICA FOR HIGHER GERMINATION PERCENTAGE AND LONGER VIABILITY

    Parveen
    Forest Research Institute, Dehradun 248006


    Introduction

    N eem (Azadirachta indica A. Juss) is a valuable multipurpose tree species for both rural and urban people. It is a versatile tree with a variety of uses in industry, medicine and timber (Radwanski and Wickens, 1981). The oil is used extensively in soap industry and the cake, the residue left after extraction of oil from the seed, makes an excellent fertilizer and soil conditioner. It is widely distributed in several parts of the world. It is found in dry areas of India, Pakistan, Sri Lanka, Malaysia, Indonesia, Thailand and Myanmar. In India, the tree is distributed in tropical dry areas up to 1,500 m altitude. It is largely found in the states of Uttar Pradesh, Haryana, Punjab, Himachal Pradesh, Rajasthan, Delhi, Gujarat, Madhya Pradesh, Bihar, Maharashtra. Andhra Pradesh, Karnataka, Tamil Nadu, Kerala, Orissa, West Bengal and Assam. It occurs naturally, but it is also planted in field and roadside plantation. It is a fast growing tree and best growth is found in areas having rainfall of about 750-1,000 mm but can survive in areas receiving only 150 mm of annual rainfall. It can tolerate long dry season, but does not tolerate excessive cold especially in the seedling and sapling stages. It coppices well, produces root suckers and withstands pollarding. The fruit of neem is an ellipsoidal drupe, about 1.25 cm long. Its colour is green and after ripening it turns yellow. The fruit is generally one seeded (rarely 2 seeded). The tree starts flowering and fruiting at age of 5 years but good yield of fruits is obtained after 10-12 years. A medium size tree produces 37-55 kg seeds (Lakshmikanthan, 1978).

    Phenological Studies

    The seeds were collected from 37 sources representing 37 districts of 11 states of India (Table 1). Mature fruits, which were yellowish and yellowish green in colour were collected in the last week of June to 3rd week of July. In some area of northern and northwest India, restricted flowering and fruiting in some trees have been observed from September to November (Dwivedi, 1993). A number of neem trees at Jodhpur were fruiting during June-July and second fruiting in September-November but only few branches bear fruits (Dwivedi, 1992). Flowering and fruiting in neem in some parts of West Bengal is reported throughout the year. Mature fruits are obtained 10-12 weeks after flowering (Maithani et al., 1989).

    Seed Processing

    Fully developed and yellowish green seeds of all seed sources were de-pulped, cleaned and dried in sun. In one experiment, seeds of Jodhpur source were de-pulped, cleaned and dried in sun and room temperature. The influence of a range of drying and storage conditions was assessed. It was found that the most efficient and early drying method was sun-drying, as compared with drying at room temperature.

    Seed Germination Studies

    The germination percentage, germination value, germination energy and energy period were calculated as per International Seed Testing Association (1976) rules (Tabel 1).

    Table 1. Seed germination studies of Azadirachta indica collected from different geographical seed sources in India

    S. no. Location State Germination
    (%)
    Germination value (%) Germination energy (%) Energy period (days)
    1 Ladpur Rajasthan 40.0 3.6 40.0 21
    2 Jaisalmer Rajasthan 75.0 25.7 55.0 8
    3 Bikaner Rajasthan 73.3 13.8 71.6 19
    4 Jodhpur Rajasthan 93.3 19.7 93.3 21
    5 Pali Rajasthan 55.8 8.5 43.8 14
    6 Sawai madhopur Rajasthan 40.0 5.5 25.0 9
    7 Sikar Rajasthan 46.6 9.9 46.6 15
    8 Satara Maharashtra 30.0 6.3 26.6 09
    9 Sholapur Maharashtra 80.0 34.3 68.8 08
    10 Pune Maharashtra 31.6 3.6 21.6 09
    11 Ravinagar Maharashtra 56.6 9.9 46.6 14
    12 Nagpur Maharashtra 5.0 0.05 5.0 22
    13 Amravati Maharashtra 20.0 1.27 18.3 16
    14 Jhansi Uttar Pradesh 53.3 10.9 33.3 09
    15 Muzzafar nagar Uttar Pradesh 68.3 16.5 56.6 13
    16 Matura Uttar Pradesh 30.3 2.2 30.0 20
    17 Kanpur Uttar Pradesh 28.3 1.6 28.3 22
    18 Palanpur Gujarat 5.0 0.1 5.0 15
    19 Gandhi nagar Gujarat 23.3 1.3 23.3 20
    20 Rajkot Gujarat 38.3 6.3 28.3 09
    21 Gurgaon Haryana 6.6 0.1 6.6 17
    22 New Delhi Delhi 40.0 4.1 26.6 13
    23 Indore Madhya Pradesh 85.0 16.3 85.0 21
    24 Ujjain Madhya Pradesh 51.6 6.0 48.3 19
    25 Bilaspur Madhya Pradesh 73.3 18.9 61.6 14
    26 Jabalpur Madhya Pradesh 1.6 0.03 1.6 09
    27 Hoshangabad Madhya Pradesh 41.6 5.4 33.3 14
    28 Shivpuri Madhya Pradesh 53.3 9.7 38.3 11
    29 Sohagi Madhya Pradesh 78.3 17.5 76.6 18
    30 Maihar Madhya Pradesh 63.3 11.9 50.0 14
    31 Katni Madhya Pradesh 3.3 0.03 3.3 19
    32 Kuthilia Madhya Pradesh 85.5 39.8 75.0 10
    33 Raipur Madhya Pradesh 55.0 8.9 43.3 14
    34 Mulug Andhra Pradesh 20.0 1.8 18.3 13
    35 Bankura West Bengal 1.6 2.7 1.6 10
    36 Ranchi Bihar 28.3 3.8 28.2 18
    37 Coimbatore Tamil Nadu 3.3 0.03 3.3 18

    Germination Percentage (GP)
    100 seeds with endocarp in three replicates were sown in germination trays filled with sand. The observation period for germinative capacity was set at 30 days.

    GP(%)= Total number of seeds germinated*100
    Total number of seeds sown

    Germination Value (GV)
    The speed of germination was expressed in terms of germination value for which Czabator’s (1962) formula was followed
    GV = (PV) × (MDG)
    PV = Highest value of cumulative germination percentage divided by number of days since start of the experiment
    MDG = mean daily germination

    Germination Energy (GE)
    Germination energy was calculated by using the following formula

    GE(%)= Number of germinating seeds during the germinative energy period*100
    Total number of seeds in the sample

    Energy Period
    This is the peak period in days which commences from the start of germination till peak period of germination

    Germination Percentage
    The highest germination was obtained in seeds from Jodhpur with 93.3 per cent germination. Seeds of Jodhpur source without endocarp were also used for germination studies. It was also found that seeds sown without endocarp require less time to start ge