ABSTRACT
Bamboo is a genetic resource with evident potential for use from construction until the recovery of degraded areas. Although, characterization and evaluation studies involving bamboo species are scarce and it is difficult to define a list of specific descriptors to better meet the different genres and or bamboo species. Thus, the objective of this work were propose and test the effectiveness of qualitative and quantitative descriptors for phenotypic characterization and the study of genetic diversity among six species of bamboo introduced in Brazil. Fifteen qualitative and nine quantitative descriptors were proposed and tested in clones with one year of planting. Individuals belonging to six species of bamboo (Bambusa vulgaris, Bambusa vulgaris var. vittata, Drepanostachyum falcatum, Dendrocalamus latiforus, Phyllostachys aurea var. albovariegada and Phyllostachys edulis) were characterized on the basis of vegetative descriptors, pseudopetiole, sheath, ligule, gems and culm. The genetic divergence between the clones was estimated by the methods of grouping of Tocher and UPGMA with use of average Euclidean distance and the principal component in two-dimensional plane. Qualitative and quantitative descriptors proposed were efficient to differentiate the six species studied and quantify genetic diversity. The quantitative descriptor of sheath length was the largest contributor to differentiate the species studied.
Key words:
Bambusa; Drepanostachyum; Dendrocalamus; Phyllostachys; Genetic diversity
RESUMO
O bambu é um recurso genético com evidente potencial para utilização que vai desde a construção civil até a recuperação de áreas degradadas. Mesmo assim, os estudos de caracterização e avaliação envolvendo espécies de bambu são escassos e dificultam a definição de uma lista de descritores específica para melhor conhecer os diferentes gêneros e/ ou espécies de bambu. Desse modo, objetivou-se neste trabalho propor e testar a eficiência de descritores qualitativos e quantitativos visando à caracterização fenotípica e ao estudo de diversidade genética entre seis espécies de bambu introduzidas no Brasil. Quinze descritores qualitativos e nove quantitativos foram propostos e testados em clones com um ano de plantio. Indivíduos pertencentes a seis espécies de bambu (Bambusa vulgaris, Bambusa vulgaris var. vittata, Drepanostachyum falcatum, Dendrocalamus latiforus, Phyllostachys aurea var. albovariegada e Phyllostachys edulis), foram caracterizados com base em descritores vegetativos de folha, pseudopecíolo, bainha, lígula, gemas e colmo. A divergência genética entre os clones foi estimada pelos métodos de agrupamento de Tocher e UPGMA, com emprego da distância Euclidiana média e pelos Componentes Principais no plano bidimensional. Os descritores qualitativos e quantitativos propostos foram eficientes para diferenciar as seis espécies estudadas e quantificar a diversidade genética. O descritor quantitativo de comprimento da bainha foi o que mais contribuiu para diferenciar as espécies estudadas.
Palavras-chave:
Bambusa; Drepanostachyum; Dendrocalamus; Phyllostachys; Diversidade genética
INTRODUCTION
The bamboo is a member of the Poaceae family, included inthe subfamily Bambusoideae, with 1,439 species (BAMBOO PHYLOGENY GROUP, 2012BAMBOO PHYLOGENY GROUP. An updated tribal and subtribal classification of the bamboos (Poaceae: Bambusoideae). The Journal of the American Bamboo Society, v. 24, n. 1, p. 1-10, 2012.; SUNGKAEW et al., 2009SUNGKAEW, S. et al. Non-monophyly of the woody bamboos (Bambuseae; Poaceae): a multi-gene region phylogenetic analysis of Bambusoideae. Journal of Plant Research, v. 122, p. 95-108, 2009.). Bamboos are distributed throughout the tropical and temperate areas of Asia, Australia, Africa and America (GUERREIRO; LIZARAZU, 2010GUERREIRO, C. I., LIZARAZU, M. A. Flowering of Bambusa tuldoides (Poaceae, Bambusoideae, Bambuseae) in southern South America. Darwiniana, v. 48, n. 1, p. 25-31, 2010.).
Brazil has a great diversity of bamboo; thirty-three genera and around 250 species can be found, of which about 160 are considered to be endemic, with some that have not yet been formally described. In addition, over 20 species of bamboo have been introduced into Brazil (FILGUEIRAS et al., 2013.FILGUEIRAS, T. S. et al. Poaceae. In: Lista de Espécies da Flora do Brasil. Jardim Botânico do Rio de Janeiro, 2013. Disponível em: http://floradobrasil.jbrj.gov.br/jabot/floradobrasil/FB102232 . Acesso em: 12 maio 2015.
http://floradobrasil.jbrj.gov.br/jabot/f...
), mainly during the colonial period and later by Japanese immigrants, most of these being species native to Asia (SILVA; PEREIRA; SILVA, 2011SILVA, I. F.; PEREIRA, D. S.; SILVA, S. R. Estudos morfológicos do bambu (Bambusa cf. vulgaris), uma espécie invasora em área de Mata Atlântica no Parque Municipal de Maceió-Alagoas. Revista Semente, v. 6, n. 6, p. 99-109, 2011.; TOMBOLATO; GRECO; PINTO, 2012TOMBOLATO, A. F. C.; GRECO, T. M.; PINTO M. M. Dez espécies de bambus exóticos mais comuns no paisagismo no Brasil. Revista Brasileira de Horticultura Ornamental, v. 18, n. 2, p. 105-114, 2012.).
There are many different uses for bamboo. Many species are grown as ornamental plants, while others are used to control erosion in degraded areas, as a source of raw material for the structure of housing, as forage, as a source of fibre for the pulp and paper industry and as a source of biomass for energy production; the edible shoots are used for food. In addition, bamboo serves as raw material for craftsmen, making it a useful resource in local economies (FILGUEIRAS et al., 2013FILGUEIRAS, T. S. et al. Poaceae. In: Lista de Espécies da Flora do Brasil. Jardim Botânico do Rio de Janeiro, 2013. Disponível em: http://floradobrasil.jbrj.gov.br/jabot/floradobrasil/FB102232 . Acesso em: 12 maio 2015.
http://floradobrasil.jbrj.gov.br/jabot/f...
; FILGUEIRAS; GONÇALVES; 2004FILGUEIRAS, T. S.; GONÇALVES, A. P. S. A checklist of the basal grasses and bamboos in Brazil (Poaceae). The Journal of the American Bamboo Society, v. 18, n. 1, p. 7-18, 2004.; RESENDE et al., 2011RESENDE, C. D. S. et al. Obtenção do extrato hidroalcoólico da Bambusa vulgaris vittata e avaliação da atividade antimicrobiana. In: SEMINÁRIO DE PESQUISAS E TCC DA FACULDADE UNIÃO DE GOYAZES, 2., 2011, Trindade,GO. Anais..., Trindade,GO, 2011. p. 1-24.).
Bamboos form a monophyletic group within the Poaceae family, and can be recognised by their morphological, anatomical, embryological, physiological, cytological, macromolecular and ecological characteristics. These characteristics are used by researchers to identify, qualify, characterise, name, classify and recommend uses for the various species that make up this rich and diverse group of plants (FILGUEIRAS; GONÇALVES, 2011FILGUEIRAS T. S.; GONÇALVES, A. P. S. Bambus nativos no Brasil: oportunidades e desafios para seu conhecimento. In: SEMINÁRIO NACIONAL DO BAMBU: estruturação da rede de pesquisa e desenvolvimento, 1., 2006, Brasília. Anais.... 2. ed. Brasília: CPAB, Universidade de Brasília, 2011. p. 196.). One of the more interesting features among bamboos is related to flowering. Flowering in most species of bamboo is irregular until the end of the vegetative stage, a stage that can last up to 120 years (AZZINI; CLARAMELLO; NAGAI, 1978AZZINI, A.; CLARAMELLO, D.; NAGAI, V. Propagação vegetativa do bambu gigante. Bragantia, v. 37, n. 1, p. 1-3, 1978.). After flowering some species die, making study of the floral characteristics of the bamboo difficult; they also produce a small number of seeds (RAMANAYAKE, 2006RAMANAYAKE, S. M. S. D. Flowering in bamboo: an enigma! Ceylon Journal of Science, v. 35, n. 2, p. 95-105, 2006.).
Most current classifications of the bamboo depend on morphological characteristics (CLARK et al., 2007CLARK, L. G. et al. Phylogenetic relationships among the one-flowered, determinate genera of Bambuseae (Poaceae: Bambusoideae). A Journal of Systematic and Evolutionary Botany, v. 23, n. 1, p. 315-332, 2007.), however few species have been properly documented, making it difficult to use phenotypic features. This is further compounded by the fact that taxonomic studies traditionally depend on inflorescence and floral morphology, since vegetative characteristics can be influenced by environmental conditions (SHALINI et al., 2013SHALINI, A. et al. Evaluation of genetic diversity in bamboo through DNA marker and study of association with morphological traits. Bulletin of Environment, Pharmacology and Life Sciences, v. 2, n. 8, p. 78-83, 2013.; YEASMIN et al., 2015YEASMIN, L. et al. Bamboo: an overview on its genetic diversity and characterization. 3 Biotech, v. 5, p. 1-11, 2015.).
Genetic diversity in the bamboo should be studied, providing data to investigate evolution of the bamboo through speciation (FILGUEIRAS; GONÇALVES, 2011FILGUEIRAS T. S.; GONÇALVES, A. P. S. Bambus nativos no Brasil: oportunidades e desafios para seu conhecimento. In: SEMINÁRIO NACIONAL DO BAMBU: estruturação da rede de pesquisa e desenvolvimento, 1., 2006, Brasília. Anais.... 2. ed. Brasília: CPAB, Universidade de Brasília, 2011. p. 196.). In addition, for bamboo to be seen as a viable alternative for its many uses, studies are necessary, so as to characterise and evaluate possible species for use in appropriate applications. However, studies into the characterisation and evaluation of bamboo species are scarce, and there is as yet no definitive list of morphological descriptors. The aim of this study therefore, was to propose and test the efficiency of qualitative and quantitative descriptors, with a view to the phenotypic characterisation and to the study of genetic diversity among six species of bamboo introduced into Brazil.
MATERIAL AND METHODS
Six accessions were used in the study, representing different species of bamboo (Bambusa vulgaris, Bambusa vulgaris var. vittata, Drepanostachyum falcatum, Dendrocalamus latiforus, Phyllostachys aurea var. albovariegada and Phyllostachys edulis). All material was provided by the Biomudas Laboratory, which maintains a collection of bamboos at Venda Nova do Imigrante, in the State of Espírito Santo (ES), Brazil. Plantlets from each species were obtained by the division method, used in the propagation of bamboo, where parts were taken from the rhizome and culm, and planted in 30-litre pots containing Basaplant® Florestais substrate.
The experiment was carried out in a greenhouse of the Research Support Unit (UAP) of the Darcy Ribeiro North Fluminense State University (UENF), at Campos dos Goytacazes, in the State of Rio de Janeiro (RJ), located at 21°45’ S and 41°20’ W at an average altitude of 11 metres.
Morphological characterisation was performed for two plants per species after around one year of growth. These were properly identified using bamboo identification keys to confirm the species (SHIRASUNA, 2012SHIRASUNA, R. T. Bambus nativos (Poaceae: Bambusoideae) no Parque Estadual das Fontes do Ipiranga, São Paulo, Brasil. 2012. 266 f. Dissertação (Mestrado em Biodiversidade Vegetal e Meio Ambiente) - Instituto de Botânica da Secretaria de Estado do Meio Ambiente, São Paulo, 2012.; WONG, 2004WONG, K. M. Bamboo: the amazing grass: a guide to the diversity and study of bamboos in Southeast Asia. Selangor Darul Ehsan, Malaysia: International Plant Genetic Resources Institute (IPGRI); Kuala Lumpur, Malaysia: University of Malaya, 2004. 80 p. Disponível em: http://www.bioversityinternational.org/ . Acesso em: 02 dez. 2013.
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).
Vegetative morphological characteristics which are noticeably variable, from such parts of the plant as the culm, branches, leaves, petioles, sheath, ligule and buds, were used to make up the proposed descriptors for the species being studied, resulting in 15 qualitative and nine quantitative characteristics (Tables 1 and 2). For the qualitative characteristics, which involved establishing the colour, a scale was used to determine colour intensity. The quantitative characteristics were measured with the aid of a digital calliper, metal tape measure and millimetre rule.
Proposed qualitative descriptors for the bamboo species: B. vulgaris, B. vulgaris var. vittata, D. falcatum, D. latiforus, P. aurea var. albovariegada and P. edulis.
Proposed quantitative descriptors for the bamboo species: B. vulgaris, B. vulgaris var. vittata, D. falcatum, D. latiforus, P. aurea var. albovariegada and P. edulis
Genetic divergence between the species, and multivariate analysis using the mean Euclidean distance as a measure of dissimilarity, were determined by Tocher clustering optimisation (RAO, 1952RAO, R. C. Advanced statistical methods in biometric research. New York: J. Wiley, 1952. 330 p.), mean hierarchical clustering between groups (UPGMA), and principal component analysis. All analyses of genetic divergence were performed using the GENES software (CRUZ, 2013CRUZ, C. D. Genes:a software package for analysis in experimental statistics and quantitative genetics. Acta Scientiarum. v. 35, n. 3, p. 271-276, 2013.).
RESULTS AND DISCUSSION
For the qualitative features, no variability was seen in the descriptors of node characteristics (NC), internode spikes (IS), shape of culm (CS), internode pilosity (IP), blade position (BP) and internode filling (IF). Therefore these descriptors did not contribute to differentiating between the species (Table 3).
Qualitative morphological characteristics in six bamboo genotypes, evaluated in a greenhouse
For sheath colouration (SC), the species B. vulgaris vitatta and D. latiforus presented light green sheaths with yellow stripes. The species P. aurea albovariegada displayed sheaths of a green colouration with white stripes. The species B. vulgaris and P edulis had medium green sheaths, and D. falcatum, light green, this being a feature that facilitated differentiation between the species. The presence of sheath wax (CSW) was only seen in P aurea albovariegada (Table 3).
The species with a culm colouration (CC) of two colours are B. vulgaris vitatta, with yellow culms with dark green stripes, and D. latiforus, with medium green culms with dark green stripes. The species P. aurea albovariegada and P. edulis had medium green culms, D. falcatum had light green culms and B. vulgaris dark green culms (Table 3).
For the descriptor of leaf-blade colour (BC), the species D. falcatum displayed clear green leaf blades, in the species B. vulgaris, B. vulgaris vitatta, latiforus D. and P edulis the blades had a dark green colouration, and in P. aurea albovariegada they were medium green with white stripes (Table 3). This characteristic for leaves of more than one colour encourages the use of the species as an ornamental plant. For the descriptor of auricle development (AD), evident auricles were seen in the species D. falcatum, B. vulgaris D. latiforus and P. edulis. At the junction of the blade and the sheath is the external ligule (LD), situated on the abaxial surface, and which may or may not be evident. Only the species B. vulgaris, P. edulis and P. aurea albovariegada displayed evident ligules (Table 3).
For the descriptor of sheath pilosity (SP), the species D. falcatum and B. vulgaris have glabrous sheaths, while those of the remaining species are pilose (Table 3).
Pilosity of the leaf blade (BPI) was evaluated for the blade (L), the ribs (R) and the borders (B). Pilosity was seen in the borders of the leaf blade in all the species under study, with the species B. vulgaris vittata and P. edulis presenting pilosity throughout the leaf-blade area. In the species D. falcatum and B. vulgaris, pilosity was seen on the blade, and in D. latiforus, pilosity was seen on the ribs (Table 3).
The quantitative characteristics contributed to differentiating between the species (Table 4).
Quantitative morphological characteristics in six bamboo genotypes, evaluated in a greenhouse
The method proposed by Singh (1981)SINGH, D. The relative importance of characters affecting genetic divergence. Indian Journal of Genetic and Plant Breeding, v. 41, n. 2, p. 237-245, 1981. made it possible to estimate the relative contribution of each quantitative descriptor to the genetic diversity between species. The quantitative characteristic that least contributed to differentiate between the species under study was seen to be plant height (PH) (0.22%). The characteristics with the greatest contribution were the length of the culm sheath (SL) (39.88%), width of the leaf (LW) (18.24%) and length of the internodes (IL) (15.4%) (Table 5).
Relative contribution of quantitative characteristics to genetic divergence between six bamboo genotypes, using a method proposed by Singh (1981)SINGH, D. The relative importance of characters affecting genetic divergence. Indian Journal of Genetic and Plant Breeding, v. 41, n. 2, p. 237-245, 1981.
Despite the low contribution of plant height to divergence, this descriptor cannot be ignored, as it is decisive when utilising the species. As pointed out by Wahab et al. (2010)WAHAB, R. et al. Anatomical and physical properties of cultivated two- and four-year-old Bambusa vulgaris. Sains Malaysiana, v. 39, n. 4, p. 571-579, 2010., plant height, number of internodes per culm, internode length and culm diameter are important factors in determining the most appropriate use for each species of bamboo.
When evaluating genetic divergence according to the values obtained for dissimilarity, the greatest genetic distance was between the species B. vulgaris vittata and P. aurea albovariegada (Table 6). The species involved belong to different genera and subtribes, which explains this result; the genus Bambusa belongs to the subtribe Bambusinae, while the genus Phyllostachys belongs to the subtribe Shibataeinae (DAS et al, 2008DAS, M. et al. Bamboo taxonomy and diversity in the era of molecular markers. Advances in Botanical Research, v. 47, p. 225-267, 2008.). The smallest genetic distance was found between the species B. vulgaris and D. latiforus (Table 6).
The Tocher method, when applied to qualitative data, allows for the formation of clusters based on the dissimilarity expressed by the divergence matrix obtained with the index ‘1-C’, where ‘C’ is the concordance of values expressed during comparison of the species. Thus, with the qualitative characteristics, the formation of three groups was seen, being a more restrictive method due to the use of discrete variables. Group 1 comprised the species D. falcatum, B. vulgaris and P. edulis. Group 2 included the species B. vulgaris vittata and D. latiforus. Finally, group 3 consisted of the species P. aurea albovariegada (Table 7).
Groups of bamboo genotypes established by the Tocher method, based on the dissimilarity expressed by the mean Euclidean distance obtained from qualitative characteristics and descriptors that facilitated the formation of groups
Four groups were formed for the quantitative characteristics. Group 1 was composed of the species B. vulgaris and D. latiforus, group 2 of D. falcatum and P. aurea albovariegada, group 3 of P. edulis and group 4 of B. vulgaris vittata (Table 8).
Groups of bamboo genotypes established by the Tocher method, based on the dissimilarity expressed by the mean Euclidean distance obtained from quantitative characteristics that most contributed to the formation of groups
The dendrogram of genetic similarity between the six bamboo genotypes, obtained by UPGMA, and based on the dissimilarity matrix of the qualitative data (Figure 1), showed that with a 40% cut it was possible to form four groups. The dendrogram based on the dissimilarity matrix of the quantitative data (Figure 2) showed that with 40% of the data, five groups were formed.
Dendrogram of the genetic similarity between six bamboo genotypes, obtained by UPGMA, and based on the dissimilarity matrix of the qualitative data
Dendrogram of the genetic similarity between six bamboo genotypes, obtained by UPGMA, and based on the dissimilarity matrix of the quantitative data
Principal component analysis reduced the nine quantitative characteristics to two principal components (PC1 and PC2), which explained 91.44% of the total variance between the six species. The greatest proximity was seen between the species D. latiforus (4) and B. vulgaris (2). The greatest distance was seen between the species P aurea albovariegada (6) and B. vulgaris vittata (3) (Figure 3).
Principal component analysis (PC1 and PC2), with 91.44% of accumulated variance, between six bamboo genotypes: (1) D. falcatum, (2) B. vulgaris, (3) B. vulgaris vittata, (4) D. latiforus, (5) P. edulis and (6) P. aurea albovariegada, based on quantitative characteristics
Based on the vegetative morphological descriptors, the species D. latiforus and B. vulgaris were seen to be very close. At the stage evaluated in this study, the two species are very similar. This was also noted by Sun, Xia and Lin (2005)SUN, Y.; XIA, N.; LIN, R. Phylogenetic analysis of Bambusa (Poaceae: Bambusoideae) based on internal transcribed spacer sequences of nuclear ribosomal DNA. Biochemical Genetics, v. 43, n. 12 p. 603-612, 2005. where, by using ribosomal DNA as molecular markers in phylogenetic analysis, the species Bambusa and Dendrocalamus were allocated to the same group. According to Li (1997)LI, D. Z. The flora of China Bambusoideae project: problems and current understanding of bamboo taxonomy in China. In:The bamboos, v. 19, p. 61-81, 1997., there are similarities between many species of the genera Bambusa and Dendrocalamus, although the relationship between these genera is still not well understood.
Classical systems for classifying bamboo are based on vegetative and reproductive morphological characteristics (DAS et al., 2007DAS, M. et al. Phylogenetic relationships among the bamboo species as revealed by morphological characters and polymorphism analyses. Biologia Plantarum, v. 51, n. 4, p. 667-672, 2007.), although the reproductive characteristics of these plants often prevent such classification. Vegetative descriptors therefore, although they may be influenced by environmental conditions, are reliable and should be used in diversity studies.
In this work, high genetic diversity was seen between the six species under study using morphological descriptors; this was also observed by Das et al. (2007)DAS, M. et al. Phylogenetic relationships among the bamboo species as revealed by morphological characters and polymorphism analyses. Biologia Plantarum, v. 51, n. 4, p. 667-672, 2007., who found high genetic diversity between 15 species of bamboo in India. Also, in studies involving different individuals of the same species, Bambusa Tulda, Bhattacharya et al. (2006)BHATTACHARYA, S. et al. Morphological and molecular characterization of Bambusa tulda with a note on flowering. Annals of Botany, v. 98, p. 529-535, 2006. found little genetic diversity as would be expected. Further studies involving a large number of bamboos from different regions are therefore required to achieve a better understanding of their genetic diversity.
CONCLUSIONS
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1. The proposed qualitative and quantitative descriptors were efficient in differentiating the species B. vulgaris, B. vulgaris var. vittata, D. falcatum, D. latiforus, P. aurea var. albovariegada and P. edulis;
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2. The species showed high genetic diversity;
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3. The largest contributor in differentiating between the species under study was the quantitative descriptor for length of sheath;
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4. The species with the greatest genetic proximity were D. latiforus and B. vulgaris. The greatest distance was seen between the species P. aurea albovariegada and B. vulgaris vittata.
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1
Parte da dissertação de mestrado do primeiro autor apresentada ao programa de pós-graduação em Genética e Melhoramento de Plantas da Universidade Estadual do Norte Fluminense Darcy Ribeiro/UENF
ACKNOWLEDGEMENTS
The authors wish to thank Capes for the grant of a masters scholarship to the lead author of this work. Thanks also go to Biomudas Laboratory for the donation of the plants used in the study.
REFERENCES
- AZZINI, A.; CLARAMELLO, D.; NAGAI, V. Propagação vegetativa do bambu gigante. Bragantia, v. 37, n. 1, p. 1-3, 1978.
- BAMBOO PHYLOGENY GROUP. An updated tribal and subtribal classification of the bamboos (Poaceae: Bambusoideae). The Journal of the American Bamboo Society, v. 24, n. 1, p. 1-10, 2012.
- BHATTACHARYA, S. et al Morphological and molecular characterization of Bambusa tulda with a note on flowering. Annals of Botany, v. 98, p. 529-535, 2006.
- CLARK, L. G. et al Phylogenetic relationships among the one-flowered, determinate genera of Bambuseae (Poaceae: Bambusoideae). A Journal of Systematic and Evolutionary Botany, v. 23, n. 1, p. 315-332, 2007.
- CRUZ, C. D. Genes:a software package for analysis in experimental statistics and quantitative genetics. Acta Scientiarum v. 35, n. 3, p. 271-276, 2013.
- DAS, M. et al Bamboo taxonomy and diversity in the era of molecular markers. Advances in Botanical Research, v. 47, p. 225-267, 2008.
- DAS, M. et al Phylogenetic relationships among the bamboo species as revealed by morphological characters and polymorphism analyses. Biologia Plantarum, v. 51, n. 4, p. 667-672, 2007.
- FILGUEIRAS T. S.; GONÇALVES, A. P. S. Bambus nativos no Brasil: oportunidades e desafios para seu conhecimento. In: SEMINÁRIO NACIONAL DO BAMBU: estruturação da rede de pesquisa e desenvolvimento, 1., 2006, Brasília. Anais... 2. ed. Brasília: CPAB, Universidade de Brasília, 2011. p. 196.
- FILGUEIRAS, T. S. et al Poaceae. In: Lista de Espécies da Flora do Brasil Jardim Botânico do Rio de Janeiro, 2013. Disponível em: http://floradobrasil.jbrj.gov.br/jabot/floradobrasil/FB102232 . Acesso em: 12 maio 2015.
» http://floradobrasil.jbrj.gov.br/jabot/floradobrasil/FB102232 - FILGUEIRAS, T. S.; GONÇALVES, A. P. S. A checklist of the basal grasses and bamboos in Brazil (Poaceae). The Journal of the American Bamboo Society, v. 18, n. 1, p. 7-18, 2004.
- GUERREIRO, C. I., LIZARAZU, M. A. Flowering of Bambusa tuldoides (Poaceae, Bambusoideae, Bambuseae) in southern South America. Darwiniana, v. 48, n. 1, p. 25-31, 2010.
- LI, D. Z. The flora of China Bambusoideae project: problems and current understanding of bamboo taxonomy in China. In:The bamboos, v. 19, p. 61-81, 1997.
- RAMANAYAKE, S. M. S. D. Flowering in bamboo: an enigma! Ceylon Journal of Science, v. 35, n. 2, p. 95-105, 2006.
- RAO, R. C. Advanced statistical methods in biometric research New York: J. Wiley, 1952. 330 p.
- RESENDE, C. D. S. et al Obtenção do extrato hidroalcoólico da Bambusa vulgaris vittata e avaliação da atividade antimicrobiana. In: SEMINÁRIO DE PESQUISAS E TCC DA FACULDADE UNIÃO DE GOYAZES, 2., 2011, Trindade,GO. Anais..., Trindade,GO, 2011. p. 1-24.
- SHALINI, A. et al Evaluation of genetic diversity in bamboo through DNA marker and study of association with morphological traits. Bulletin of Environment, Pharmacology and Life Sciences, v. 2, n. 8, p. 78-83, 2013.
- SHIRASUNA, R. T. Bambus nativos (Poaceae: Bambusoideae) no Parque Estadual das Fontes do Ipiranga, São Paulo, Brasil 2012. 266 f. Dissertação (Mestrado em Biodiversidade Vegetal e Meio Ambiente) - Instituto de Botânica da Secretaria de Estado do Meio Ambiente, São Paulo, 2012.
- SILVA, I. F.; PEREIRA, D. S.; SILVA, S. R. Estudos morfológicos do bambu (Bambusa cf. vulgaris), uma espécie invasora em área de Mata Atlântica no Parque Municipal de Maceió-Alagoas. Revista Semente, v. 6, n. 6, p. 99-109, 2011.
- SINGH, D. The relative importance of characters affecting genetic divergence. Indian Journal of Genetic and Plant Breeding, v. 41, n. 2, p. 237-245, 1981.
- SUN, Y.; XIA, N.; LIN, R. Phylogenetic analysis of Bambusa (Poaceae: Bambusoideae) based on internal transcribed spacer sequences of nuclear ribosomal DNA. Biochemical Genetics, v. 43, n. 12 p. 603-612, 2005.
- SUNGKAEW, S. et al Non-monophyly of the woody bamboos (Bambuseae; Poaceae): a multi-gene region phylogenetic analysis of Bambusoideae. Journal of Plant Research, v. 122, p. 95-108, 2009.
- TOMBOLATO, A. F. C.; GRECO, T. M.; PINTO M. M. Dez espécies de bambus exóticos mais comuns no paisagismo no Brasil. Revista Brasileira de Horticultura Ornamental, v. 18, n. 2, p. 105-114, 2012.
- WAHAB, R. et al Anatomical and physical properties of cultivated two- and four-year-old Bambusa vulgaris Sains Malaysiana, v. 39, n. 4, p. 571-579, 2010.
- WONG, K. M. Bamboo: the amazing grass: a guide to the diversity and study of bamboos in Southeast Asia. Selangor Darul Ehsan, Malaysia: International Plant Genetic Resources Institute (IPGRI); Kuala Lumpur, Malaysia: University of Malaya, 2004. 80 p. Disponível em: http://www.bioversityinternational.org/ . Acesso em: 02 dez. 2013.
» http://www.bioversityinternational.org/ - YEASMIN, L. et al Bamboo: an overview on its genetic diversity and characterization. 3 Biotech, v. 5, p. 1-11, 2015.
Publication Dates
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Publication in this collection
Mar 2016
History
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Received
01 Feb 2014 -
Accepted
07 Oct 2015