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Genetic diversity of woody bamboos - their conservation and improvement - A.N. Rao

Consultant, International Plant Genetic Resources Institute (IPGRI), Serdang, Selangor Darul Ehsan, Malaysia.

Introduction

Bamboos are very interesting plants in their growth, morphogenesis, taxonomy, distribution, ecology and reproduction. The monopodial or sympodial habit, well-segmented culms and rooting patterns characterize various growth forms of bamboos. The functioning of intercalary meristem that supports rapid growth of internodes and their elongation to establish the erect stem axis that reaches 30-40 m in certain species, characterizes the morphogenesis of bamboo plants. The perennial woody habit of bamboos separates them from herbaceous grass genera. The interrelationship between various bamboo genera and species is not well understood. The complicated life history and widely variable vegetative characters create impediments in solving the taxonomic problems.

The grass family Gramineae is one of the biggest of the five among flowering plants, the tribe bambuseae includes 85% of woody and 25% herbaceous genera. Of these about 10 genera and 30 species are commercially important worldwide. In the priority list, prepared in 1993, nine genera and nineteen species were included which indicate the fact that traditionally only a small part of bamboo resources are used to date (Williams and Rao 1994; Rao et al. 1998). The scientific details available on bamboos are inadequate, except taxonomic categorization, to promote work on genetic analysis and conservation.

Regarding distribution, most of the woody genera are present in South and South East Asia which include Yunnan province in China bordering the lower Himalayas that extend to the east (Tewari 1992; Pei 1997; Rao et al. 1997; Rao et al. 1998). Ecology of bamboos is diverse in terms of habitats, soil, altitudes, water requirements, species association and others. Most of them are rapid colonisers. Vegetative growth is varied even between two species belonging to same genus. Gregarious or sporadic flowering with long or short interval of time with varying degrees of sterility would characterize sexual reproduction (Tewari 1992). No correlation has been observed between vegetative and reproductive growth phases. The clumps of woody bamboos can grow up to 100 or 120 years and each culm can last 4-10 years.

The present state of knowledge on some of the important topics that help to understand the genetic variations, conservation and production of woody bamboos are discussed and lines of research that need to be pursued in the near future are briefly specified in this paper.

Taxonomic status

Gramineae, or the grass family, has 14 tribes and Bambuseae is one of them that includes bamboos. Seedling characters, leaf anatomy, number as well as morphology of chromosomes of panicoid, chloridoid, festucoid and bambusoid members were compared to assess the evolutionary trends. The bambuseae is regarded as the early or primitive tribe among all grasses (Stebbins 1950, 1971). This generalization was made on the data obtained from temperate grasses and the criteria 'established cannot be uncritically followed' with reference to all the species, especially when the groups are distantly related to one another (Stebbins 1971). It was suggested that the data obtained indicate the need for the revision of various tribes of the whole family (Stebbins 1971). It is not certain whether bambusoid grasses examined by the above authors were of woody or herbaceous groups. A normal or standardized bamboo classification system, which is easy to use, is very much needed consolidating all the criteria so far used by different researchers in various parts of the world (McClure 1966; Ohrenberger and Goerrings 1989; Tewari 1992; Dransfield and Widjaja 1995). Taxonomic problems in species identification are common to all the bamboo growing countries in Asia with the difference that in some countries few species remain yet to be identified while in others more than 30-40% of the species remain unidentified. Further, the variations that may be present within the same species that has very wide or Pan Asian distribution have yet to be accounted and recorded (E.g. Bambusa bambos, Dendrocalamus strictus and others), because no individual scientist has studied the populations of the same species in different countries or geographical areas.

Nearly 10-20% of bamboo species that have been used for ages have no scientific names. Herbarium specimens are inadequate in many Asian countries in evaluating all the diagnostic characters. The living plants show many more useful characters. Such details are illustrated with reference to bamboos in Southeast Asia (Chua et al. 1996; Dransfield and Widjaja 1995). Nevertheless the extraction, use and trade of bamboos continue in all the countries using common and local names. Some species in certain countries are identified only with numbers.

Up to 90% of the sympodial bamboos presently used are extracted directly from the forests. Resources are dwindling. Very few countries have made any efforts so far to assess the bamboo resources available in terms of forest land area covered with bamboos, all the species present, the quality and quantity of species distributed etc (Fu and Xiao 1996; Bennet and Gaur 1990; Anon 1995, 1997; Rao et al. 1987; Rao et al. 1990).

Species delimitations in bamboos are determined by using a number of vegetative and reproductive characters and their coefficients. Since bamboos flower irregularly and at very long intervals it is not easy to study the flowering materials hence both vegetative and floral characters are used simultaneously (Holltum 1956, 1958; Gilliland 1971; Tewari 1992). The difficulties encountered in taxonomic studies on bamboos are well summarized and the basic frame work laid down 130 years ago by pioneers remains still valid, continuously used by others with or without additional details. (McClure 1966; Tewari 1992). A number of new species and genera are added in the meanwhile (Soderstrom and Ellis 1987; Dransfield and Widjaja 1995). More research is needed by native taxonomists working in bamboo growing countries so that they will have an opportunity to study the growing plants regularly and use fresh materials rather than studying specimens in herbaria. Occasionally they may be able to collect flowering specimens, which would be an additional advantage. The continued studies in unexplored areas would help to add more new taxa to the group.

Literature on bamboo taxonomy focuses on delimitation of genera and species and various workers have emphasized the need to study large number of species to regroup them and to streamline the phylogenetic relationships (Tewari 1992). The 'critical taxonomic work' on Southeast Asian bamboos is progressing (Dransfield and Widjaja 1995). Complete inventory of species in each bamboo growing country, the patterns of distribution and population diversity, the relative abundance with reference to ecological conditions should be properly recorded for better conservation and management of bamboo resources. For detailed ecogeographic survey and complete inventory selected locations should be identified in each country and sampling strategies with particular details relevant to each country should be outlined. The ecological characters including soil quality and water availability should be outlined. The associated plant species that grow with bamboos in a given habitat that help the ecosystem functioning need to be properly documented to prepare plans for conservation methods.

Flowering and seed production

McClure (1966) reviewed flowering behaviour of bamboos identifying species that are permanently sterile (E.g. Bambusa vulgaris); and those that regularly flower (E.g. Bambusa lineata, Ochlandra stridula, Schizostachyum spp). Other species fall in between the two boundaries showing monocarpic tendency irrespective of the fact that they flower sporadically or gregariously and of the interval between the two flowering periods. Many species of Phyllostachys and Arundinaria do not die after flowering. McClure also suggested that life cycles and flowering cycles of various bamboo species should be properly studied among various populations in different countries.

Flowering is infrequent, that too at long intervals, in most bamboo species and some of them grow up to 100-120 years before they start flowering. Monocarpic flowering habit in bamboos is a relic tendency and appears to be a carried over phenomenon resembling other grasses. Certain species became perennials with pronounced woody habit but monocarpic flowering habit remained as a surviving evolutionary trace. Many interesting articles have been written on flowering in bamboos (Jansen 1976; Liese 1985; Dwivedi 1990; Tewari 1992) and three main types of flowering recognised: a) Gragarious flowering occurs in populations very synchronously lasting over a period of 2-3 years; b) Sporadic flowering observed in some plants within the populations; sometimes very regular and seeds collected annually as observed in some species; whether the same plant continues to grow further even after flowering or dies has to be confirmed in many species. The details available at present are not precise (McClure 1966; Tewari 1992); c) Species of Schizostachyum and other bamboos flower regularly with or without seeds and the same plant continues to grow year after year.

Sporadic flowering is regular in bamboo species that grow in Kanchanaburi area of Thailand (about 120 miles from Bangkok). The temperature, rainfall and soil conditions appear to be most suitable since many species of about six genera both in the bambusetum and in the nearby forest flower regularly. Clump after clump in a population take turn to flower. Seeds are collected either once or twice every year. Although many hypotheses were put forward in the past, it is becoming increasingly clear in the last 5-10 years that sporadic flowering is very common in many species as observed in different countries. In a recent trip to Myanmar the author along with Dr Nyan Htun noticed several (1-2%) clumps of Bambusa blumeana in full flowering but with few seeds. Such observations need to be properly confirmed, collated and published (Figs 1-5). It is also necessary to study the populations more regularly in different localities every year and record the data. Further, plants from such populations need to be collected and grown in a suitable place to establish and conserve the gene pool (Banik 1995). Such an exercise would help to determine whether the causal factors that induce flowering are ecological or genetic in nature besides providing a reliable collection of germplasm for seed collection. Phytotron conditions or even large growth chambers may be suitable for experimentation. Because of the huge size of bamboos plants and lack of secondary cambium, it may be difficult to grow them in growth chambers. This point would bring in other questions like how much would culm or rhizome contribute towards flowering? The bamboo clumps that flower under natural conditions remain almost leafless. What is the role of foliage in inducing flowering? Why should foliage wait to develop till axis development is complete? Role of auxiliary buds etc? All these details need to be well studied to understand the causal factors and relationship between vegetative growth and flowering in bamboos.

It is generally observed that many species with gregarious flowering would produce no viable seeds. Very few studies are done on percentage fruit set, seed viability, germination etc. In the sporadic flowering types, the ratio between fertile and infertile seeds vary between the clumps or different clumps in a population. Seed sterility involves many factors including pollination, relative stage of embryo development, relationship between endosperm and embryo, thickness of seed coat and others. There are very few studies on development of bamboo seeds since the material is almost unobtainable or available. Bamboo growers usually soak the seeds before germinating them in the nursery. Only those that sink are used and the ones that float are rejected with the assumption that the latter are infertile with or without regular seed contents. Work on seed anatomy is required. The genetic implications in seed development are several which are yet to be properly analyzed. All such details play an important role while selecting superior populations for in situ or ex situ conservation.

Polyploidy and hybridization in bamboos

Polyploidy is more common among plants than in animals. More than 35% of flowering plants are polyploids (Stebbins 1971; Heywood 1995). Highest percentage of polyploids recorded are perennial herbs and bamboos and they are mostly self incompatible as recorded in certain bamboo species. It is suggested that the rhizomatous perennial habit drains the photosynthates to build up the vegetative biomass suppressing or postponing the event of flowering until the end of vegetative growth period as seen in many of the monocarpic plants including bananas, some palms and bamboos. The period recorded for completion of vegetative growth before flowering is the longest in bamboos among all the angiosperms. The period also varies among different species from a few decades to more than a century.

In polyploids the sexual reproduction system is modified or upset involving apomictic and parthenocarpic tendencies and substitution of asexual reproduction for sexual; the common examples are grasses. Loss of fertility and seed production is another important character of polyploids, commonly noticed in certain bamboo species. Chromosome segregation during mitosis or meiosis is not studied among bamboos. The autopolyploids and allopolyploids are identified by studying and matching chromosomes at metaphase in mitosis and these details are wanting for bamboos. Most of the woody bamboos so far studied are polyploids and diploids are rarely found in any of them (Table 1).

Genetic diversity within the species is the main building block for evolution and speciation. Both the genetic and species divergence have yet to be well studied among bamboos. Just in one generation the polyploids become separate and distinct from the diploid species. The concept of species is still being debated (Heywood 1995). Bamboos are such a diverse group and provide many good examples to analyze and discuss the morphological, ecological and genetic concepts of species. The number of species, their geographic range of distribution, species and ecosystem diversity are important to determine the in situ conservation programme and select the appropriate species from good populations for ex situ conservation. Many of the above details are yet to be well studied in case of tropical woody bamboos. Population studies would help to plan the conservation activities. The range of geographic distribution for certain bamboos species are recorded (Dransfield and Widjaja 1995; Rao et al. 1998).

Table 1. Polyploidy in bamboos

Source: Tewari 1992, Darlington and Janaki Ammal 1945, Darlington and Wiley, 1955).

Genus and number of species studied

Chromosome numbers in different species

1. Arundinaria 14 sps

4 species with 24 chromosomes, 8 species with 48, one with 54, one with 70 or 74 =14

2. Bambusa 18 sps

1 species with 70 or 74, 1 with 70 or 72, 3 with 64-72, 3 with 64, 2 with 48, 4 with 72, 1 with 52 or 64, 2 species with 56 or 64 or 72, 1 with 96 = 18

3. Dendrocalamus 8 sps

7 with 72, 1 with 48, or 64 or 72 = 8

4. Phyllostachys 21 sps

18 with 48, 1 with 44 or 48, 1 with 48 or 54, 1 with 70 or 74 = 21

(Haploid chromosome numbers 2, 8, 6 or 12 but not certain, somatic numbers vary from 48-96).
Depending on the evolutionary tendencies hybridization is possible among polyploids between closely or widely related species or even among different genera (Stebbins 1971). Such details are recorded in Bromus and the other grass species. The same proclivity is observed among bamboo genera and species. Interspecific hybrids were obtained from among species of Bambusa, intergeneric hybrids between Bambusa and Dendrocalamus sps. Phyllostachys and Dendrocalamus and others (Table 2). (Fu 1995; Zhang 1986, 1987, 1997). Superior bamboo hybrids with good vitality, reproductive potential and adaptability are cultivated in several provinces in China, covering more than 600 ha. The growth rate of culms, culm form, fibre length, resistance to cold (up to -5°C) of these hybrids were analyzed. More research in this direction holds great promise for the possible genetic enhancement of the species or groups of species concerned. Some of the hybrid bamboos are already introduced to other countries outside China.

Table 2. Cross-breeding between different genera and species of bamboo

Female parent Male parent

Dendrocalamus latiflorus

Sinocalamus minor

Bambusa textilis

Bambusa pervariabilis

Bambusa sinospinosa

Bambusa chungii

Phyllostachys edulis

D. latiflorus


*

*

*

X


X

S. minor








B. textilis

X



X



X

B. pervariabilis

X


X




X

B. sinospinosa

X


X





B. chungii








P. edulis



X

X

X

X


D. latiflorus +




*




B. textilis








D. latiflorus +




X




P. edulis








D. latiflorus +



X





B. pervariabilis








P. edulis +





X



S. minor








Note: * means excellent crossbreeding combination, good hybrids obtained (Fu 1995)
Biodiversity and genetic diversity

The word biodiversity came into usage in 1986 and the three levels of biodiversity namely, genetic diversity, species diversity and ecological diversity are yet to be properly investigated in bamboos (Tewari 1992). But no data are so far available on the biodiversity or genetic diversity or genetics of population of any species of bamboos. Biodiversity and sustainability are inseparable, said to be the two sides of the same coin (Wilson 1988; Reaka Kudla et al. 1997; Raven 1998). Genetic diversity attributes to the genes and genetic make up of individuals within the species. The meaning and implications of both bio and genetic diversity of bamboos were recently discussed (McNeely 1996; Stapleton and Ramanatha Rao 1996). Details published on species diversity are very few and intraspecific details recorded are few. A large number of taxa need to be studied to establish inter and intraspecific relationships. Cladistic relationships are built to determine phylogenetic affinities and DNA techniques have been used (Watanabe et al. 1994). Strategies for bamboo improvement were outlined (Banik and Rao 1996). Population studies are important to analyze the diversity within and between species. Proper guidelines should be developed for local and regional action. Ecological priority actions are needed (Vivekanandan et al. 1998; Wan Razali et al. 1994).

Spontaneous mutants were detected showing morphological variations in colour, shape and structure in Bambusa vulgaris, B. bambos, Oxytenanthera abyssinica, Phyllostachys edulis and Guadua angustifolia (Venkatesh 1992). Both intra and inter clump variations were common among these species and most of them are propagated vegetatively. Greater can be the variations to select superior plants if seeds are used as planting materials but seeds are rarely formed in the above species. Genetic variations are recorded in certain species due to outcrossing, segregation and recombination (Zhang and Chen 1987).

Intra and inter clump variations in Gigantochloa scortechinii were analyzed using electrophoretic methods (Ratnam et al. 1994). Lateral buds from 30-40 culms of three clumps were analysed using eleven enzymes. Variations were observed in three enzyme systems, when buds from different clumps or buds on the same culm were studied indicating the possible somatic mutation in the species (Ratnam et al. 1994). G. scortechinii is well naturalized in the logged over forests of Peninsular Malaysia. It is a very useful bamboo for handicrafts, basket making and other small-scale industries. Attempts are also made to establish plantations for which improved planting materials are needed (Othman and Nor 1994; Azmy 1994). Electrophoretic methods were used to determine both intra and interclump variations as well as somatic mutations. Only a small sample size was used and 3.3% frequency of somatic mutation was recorded. A large number of samples need to be tested to determine the genetic diversity and to select superior planting materials (Ratnam et al. 1994). Seeds of Bambusa bambos and Dendrocalamus strictus responded well when treated with colchicine showing increase in plant size and greater vigour (Ueda 1962; Tewari 1992). Both cytological and genetic research should progress further to identify the factors involved to improve the quality of bamboos

Similarity of structures in closely related species is called Homology, a biological concept more frequently used while describing animals than plants. The unique characters that bamboos display show strong tendencies of homology, especially in relation to culm sheath, arrangement of axillary buds, leaf arrangement, branching and others. Some of these characters are very identical in the closely related bamboo species and one has to be very critical in distinguishing and differentiating the variations among the characters that help to prepare the taxonomic keys and identify the species. The genetic basis of homology is not well established in plants to explain inter and intra specific relationships (Tautz 1998). A very close study of developmental characteristics in bamboo species will be very interesting to understand the role of ecological and genetical factors that determine the genotypic and phenotypic characters.

Conclusions

The work on genetic diversity, patterns of distribution, and herbarium survey of bamboos done in the last 5-6 years has given us better insight to continue research on conservation of important bamboos including the priority species. Concepts of biodiversity and sustainable use are very closely intertwined, not easy to separate or study one aspect without referring to the other. Relative evolutionary history of bamboos, endemism, geology and characterization of habitat are all inter-related and their implications need to be analyzed to plan and carry out research on methods of conservation. Evolutionary tendencies are not properly traced or determined in bamboos and information obtained on these lines would help to strengthen taxonomic relationships of bamboos and to identify the advanced, superior taxa. There are no fossil evidences of bamboos. Sexuality and its function in plants is an important strategy to generate genetic variation but many bamboos do not have a regular reproductive cycle. More critical studies are needed to analyze the implications of the present reproductive methods known in bamboos including absence of seed set, ineffective pollination, fertilization and other related aspects. It is said that economy is a subsidiary of the environment and restoration ecology, use of degraded and waste lands are important to grow bamboos that would help regenerate plant wealth and improve the economy of rural people in Asian countries who are the main bamboo users. Studies on the lines indicated above would provide good perspective for better management of superior bamboos including their conservation and sustainable use.

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Fig. 1. Bambusa blumeana, vegetative and reproductive phases.

Fig. 2. Bambusa blumeana, vegetative and reproductive phases.

Flowering and fruiting in bamboo

Fig. 3. Dendrocalamus hamiltonii

Fig. 4. Dendrocalamus sp.

Fig. 5. Dendrocalamus sp.


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