Anatomia, densidade e condutividade hidráulica potencial do xilema secundário de árvores de três procedências de Balfourodendron riedelianum (Engl.) Engl. (Rutaceae) em plantios homogêneos
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Data
2018-02-28
Autores
Silva, Jane Rodrigues da
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Universidade Estadual Paulista (Unesp)
Resumo
Estudos intraespecíficos com procedências replicadas em diferentes locais permitem distinguir o efeito do genótipo e do ambiente no crescimento e na estrutura anatômica das plantas. Diante das previsões de intensificação dos períodos seca e o aumento da temperatura em todo o mundo, entender como a predisposição genética e o ambiente influenciam o crescimento das plantas e as características do xilema secundário é importante para estabelecer futuros padrões na distribuição de espécies arbóreas. Assim, neste trabalho, avaliamos o efeito da procedência e das condições ambientais na variabilidade intraespecífica do crescimento das árvores e da estrutura anatômica do xilema secundário de Balfourodendron riedelianum (Engl.) Engl. (Rutaceae). Comparamos o crescimento (estimado por meio da altura e do diâmetro do caule), a densidade da madeira, a condutividade hidráulica potencial e as características anatômicas do xilema secundário de árvores adultas de três procedências de B. riedelianum crescidas em dois plantios homogêneos como testes de procedências. Selecionamos 72 árvores de 30 anos de idade de B. riedelianum crescidas em um plantio homogêneo localizado na Estação Experimental de Luís Antônio e outro na Estação Experimental de Pederneiras, ambos no estado de São Paulo. O solo do plantio de Luís Antônio é argiloso e neste local há maior precipitação anual do que no plantio de Pederneiras, onde o solo é arenoso. As procedências vêm de sementes coletadas em populações naturais localizadas no município de Alvorada do Sul, estado do Paraná, e Bauru e Gália, estado de São Paulo. Em Alvorada do Sul e Gália, as sementes foram coletadas em área de Floresta Estacional Semidecídua, e em Bauru foram coletadas em área de ecótono entre Cerrado e Floresta Estacional Semidecídua. Nas regiões de Alvorada do Sul e Gália há maior precipitação anual do que na região de Bauru. Para o estudo anatômico do xilema secundário, processamos o material de acordo com as técnicas usuais em anatomia do xilema secundário. Para cada indivíduo amostrado calculamos a condutividade hidráulica potencial e mensuramos a densidade da madeira e 21 características anatômicas do xilema secundário. Para testar o efeito da procedência, dos fatores ambientais, da interação entre procedência e o ambiente e a altura das árvores nas características do xilema secundário realizamos análise de variância multivariada permutacional (PERMANOVA). Para verificar as diferenças no crescimento das árvores e nas características do xilema secundário entre as procedências e entre os plantios realizamos uma análise de variância (ANOVA) de dois fatores. Identificamos que as variações nas características do xilema secundário de B. riedelianum são determinadas pela procedência, com efeito das condições ambientais dos plantios, e pela interação das procedências com as condições ambientais dos plantios. Encontramos vasos mais estreitos e com paredes celulares finas, mais agrupados e em maior densidade, com pontoações intervasculares menores e aberturas estreitas, raios mais baixos e numerosos e menor condutividade hidráulica potencial, características que indicam maior segurança no transporte de água nas árvores oriundas de Bauru, local com maior restrição hídrica, quando comparamos aos indivíduos oriundos de Gália, local com maior disponibilidade hídrica. Entretanto, para a procedência de Alvorada do Sul, apesar de ser local com condições ambientais mésicas, as árvores apresentaram xilema secundário com características que indicam maior segurança no transporte de água. Estes resultados demonstram que as procedências de Bauru e Gália mantiveram algumas características anatômicas do xilema secundário relacionadas ao seu local de origem, independente de seu local de crescimento. As condições ambientais dos plantios tiveram efeito no crescimento das plantas e nas características do xilema secundário. Encontramos árvores mais altas e com caules mais largos e o xilema secundário com características de maior eficiência no transporte de água, com maior condutividade hidráulica potencial, vasos mais largos, elementos de vaso longos, pontoações intervasculares maiores e com abertura largas e raios menos numerosos no plantio de Luís Antônio, local com maior disponibilidade de água. Isto indica que o crescimento das árvores e as características do xilema secundário de B. riedelianum também apresentam plasticidade às novas condições ambientais de crescimento.
Intraspecific studies with provenances replicated at different sites allow to distinguish the effect of genotype and environment on the growth and anatomical structure of plants. Considering predictions of drought intensification and increased temperature in wordwilde, understanding how genetic predisposition and environment influence plant growth and secondary xylem features is important to establishing future patterns in the distribution of tree species. Thus, in this work, we evaluated the effect of the provenance and environmental conditions on the intraspecific variability of tree growth and the anatomical structure of the secondary xylem of Balfourodendron riedelianum (Engl.) Engl. (Rutaceae). We compared growth (estimated from stem height and diameter), wood density, potential hydraulic conductivity, and anatomical features of the secondary xylem of adult trees of three provenances of B. riedelianum from two homogeneous plantations as provenances tests. We selected 72 trees of 30 years old of B. riedelianum grown in a homogeneous plantation located in the Experimental Station of Luís Antônio and another in the Experimental Station of Pederneiras, in the state of São Paulo. The plantations of Luís Antônio have clay soil and highest annual precipitation than the plantations of Pederneiras, where the soil is sandy. The provenances came from seeds collected in natural populations located in the city of Alvorada do Sul, state of Paraná, and from Bauru and Gália, state of São Paulo. In Alvorada do Sul and Gália, the seeds were collected in a Semideciduous Seasonal Forest, and in Bauru were collected in an ecotone between Cerrado and Semideciduous Seasonal Forest. In the regions of Alvorada do Sul and Gália there is higher annual precipitation than in the region of Bauru. For the anatomical study of the secondary xylem, we process the material according to the usual techniques in wood anatomy. For each individual sample we calculated the potential hydraulic conductivity and measured the wood density and 21 anatomical features of the secondary xylem. To test the effect of provenance, environmental factors, interaction between provenance and environmental, and tree height in the secondary xylem features, we performed permutational multivariate analyses of variance (PERMANOVA). To verify the differences between the growth and traits of the secondary xylem between the provenances and the environmental conditions, we performed a two-way ANOVA. We identified that the variations in the features of the secondary xylem of B. riedelianum are determined by the provenance, with effect of the environmental conditions of the plantations, and by the interaction of the provenances with the environmental conditions. We found narrower vessels with thinner cell walls, more grouped vessels and higher density, with smaller intervessel pits and narrow aperture, lower and numerous rays and lower potential hydraulic conductivity, features that indicate safety of water transport in the trees from Bauru, site with lower water availability, when compared to individuals from Gália, site with higher water availability. However, for provenance of Alvorada do Sul, despite being local with mesic environments, trees have secondary xylem with features that indicate safety of water transport. These results demonstrate that the provenances of Bauru and Gália maintained some anatomical features of the secondary xylem related to their site of origin, independent of their growth site. The environmental conditions of the plantations had an effect on the plants growth and secondary xylem features. We found taller trees with larger stems and secondary xylem with characteristics of higher efficiency in water transport, with higher potential hydraulic conductivity, larger vessels, longer vessel elements, larger intervessel pits with larger aperture and less number of rays in the Luís Antônio plantations, site with higher water availability. This indicates that the growth and secondary xylem features of B. riedelianum also present plasticity to the new environmental conditions of growth.
Intraspecific studies with provenances replicated at different sites allow to distinguish the effect of genotype and environment on the growth and anatomical structure of plants. Considering predictions of drought intensification and increased temperature in wordwilde, understanding how genetic predisposition and environment influence plant growth and secondary xylem features is important to establishing future patterns in the distribution of tree species. Thus, in this work, we evaluated the effect of the provenance and environmental conditions on the intraspecific variability of tree growth and the anatomical structure of the secondary xylem of Balfourodendron riedelianum (Engl.) Engl. (Rutaceae). We compared growth (estimated from stem height and diameter), wood density, potential hydraulic conductivity, and anatomical features of the secondary xylem of adult trees of three provenances of B. riedelianum from two homogeneous plantations as provenances tests. We selected 72 trees of 30 years old of B. riedelianum grown in a homogeneous plantation located in the Experimental Station of Luís Antônio and another in the Experimental Station of Pederneiras, in the state of São Paulo. The plantations of Luís Antônio have clay soil and highest annual precipitation than the plantations of Pederneiras, where the soil is sandy. The provenances came from seeds collected in natural populations located in the city of Alvorada do Sul, state of Paraná, and from Bauru and Gália, state of São Paulo. In Alvorada do Sul and Gália, the seeds were collected in a Semideciduous Seasonal Forest, and in Bauru were collected in an ecotone between Cerrado and Semideciduous Seasonal Forest. In the regions of Alvorada do Sul and Gália there is higher annual precipitation than in the region of Bauru. For the anatomical study of the secondary xylem, we process the material according to the usual techniques in wood anatomy. For each individual sample we calculated the potential hydraulic conductivity and measured the wood density and 21 anatomical features of the secondary xylem. To test the effect of provenance, environmental factors, interaction between provenance and environmental, and tree height in the secondary xylem features, we performed permutational multivariate analyses of variance (PERMANOVA). To verify the differences between the growth and traits of the secondary xylem between the provenances and the environmental conditions, we performed a two-way ANOVA. We identified that the variations in the features of the secondary xylem of B. riedelianum are determined by the provenance, with effect of the environmental conditions of the plantations, and by the interaction of the provenances with the environmental conditions. We found narrower vessels with thinner cell walls, more grouped vessels and higher density, with smaller intervessel pits and narrow aperture, lower and numerous rays and lower potential hydraulic conductivity, features that indicate safety of water transport in the trees from Bauru, site with lower water availability, when compared to individuals from Gália, site with higher water availability. However, for provenance of Alvorada do Sul, despite being local with mesic environments, trees have secondary xylem with features that indicate safety of water transport. These results demonstrate that the provenances of Bauru and Gália maintained some anatomical features of the secondary xylem related to their site of origin, independent of their growth site. The environmental conditions of the plantations had an effect on the plants growth and secondary xylem features. We found taller trees with larger stems and secondary xylem with characteristics of higher efficiency in water transport, with higher potential hydraulic conductivity, larger vessels, longer vessel elements, larger intervessel pits with larger aperture and less number of rays in the Luís Antônio plantations, site with higher water availability. This indicates that the growth and secondary xylem features of B. riedelianum also present plasticity to the new environmental conditions of growth.
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genótipo, mudanças climáticas, plasticidade fenotípica, predisposição genética, teste de procedência, genotype, climatic changes, phenotypic plasticity, genetic predisposition, provenance test