Dry semideciduous and deciduous forests occur only on calcium-rich soils and occupy almost 20% of the 200 million ha of the cerrado region of central Brazil. Other savanna physiognomies of woodlands and grasslands, and gallery forests generally occur on acid soils with low calcium levels. The literature on phytosociological aspects of such cerrado physiognomies is quite abundant whereas there is very little information on deciduous forests. Also lacking is information on soil fertility and its influence on species distribution. The objective of the present study was to study the distribution of native species within a dry deciduous forest in the Araguari river valley, Uberlâ ndia, Minas Gerais as related to soil properties. A 1-ha sampling area, with no apparent difference in physiognomy, was divided into fifty 10 m · 20 m plots arrayed in 10 · 5 grid and all trees with a minimum circumference of 15 cm at 1.30 m height were surveyed. A composite soil sample was collected from 0 to 10 cm depth in each plot and soil pH, organic C and nutrient availability were determined. Cluster analysis based on soil chemical properties showed the existence of two distinct groups of plots. Further field examinations revealed that the forest was situated in an area of transition between mica schist and basalt parent materials, without any apparent difference in the forest physiognomy. The basalt-derived soil in 23 plots had significantly higher availability of Ca (47.0 cmol(+) kg -1 soil), Mg (2.59), and K (0.96) than the mica schist derived soil (Ca-18.4, Mg-1.29, and K-0.66) in the remaining 27 plots, though both soils would be classified as eutrophic. A total of 59 species belonging to 27 families were encountered in the area as a whole, of which 36 were common to both soils. In total, 16 species occurred exclusively on the mica schist soil and 7 on the basaltic soil. Myracrodruon urundeuva Allem. (Anacardiaceae) and Tabebuia roseo-alba (Ridl.) Sandwith (Bignoniaceae) were species with high importance values in the forest as a whole. Though there was no significant difference in the diversity index between the two soils, the importance values of these species was larger in the basalt-derived soil with higher nutrient availability.Joã o Paulo De Souza is a scholarship holder of CNPq.
The rise in atmospheric CO2 concentration ([CO2]) has been accompanied by changes in other environmental factors of global climate change, such as drought. Tracking the early growth of plants under changing conditions can determine their ecophysiological adjustments and the consequences for ecosystem functions. This study investigated long-term ecophysiological responses in three woody Cerrado species: Hymenaea stigonocarpa Mart. ex Hayne, Solanum lycocarpum A. St.-Hil. and Tabebuia aurea (Silva Manso) Benth. and Hook. f. ex S. Moore, grown under ambient and elevated [CO2]. Plants were grown for 515 days at ambient (430 mg dm–3) or elevated [CO2] (700 mg dm–3). Some plants were also subjected to water stress to investigate the synergy between atmospheric [CO2] and soil water availability, and its effect on plant growth. All three species showed an increase in maximum net photosynthesis (PN) and chlorophyll index under high [CO2]. Transpiration decreased in some species under high [CO2] despite daily watering and a corresponding increase in water use efficiency was observed. Plants grown under elevated [CO2] and watered daily had greater leaf area and total biomass production than plants under water stress and ambient [CO2]. The high chlorophyll and PN in cerrado plants grown under elevated [CO2] are an investment in light use and capture and higher Rubisco carboxylation rate, respectively. The elevated [CO2] had a positive influence on biomass accumulation in the cerrado species we studied, as predicted for plants under high [CO2]. So, even with water stress, Cerrado species under elevated [CO2] had better growth.
Growth and leaf nutrient content were compared in young potted plants of Copaifera langsdorffii in sunny and shaded areas without water stress. Besides, carbon assimilation and leaf water relations were evaluated by net photosynthesis, potential photochemical efficiency and leaf water potential during daily courses in dry and rainy periods under natural conditions in both contrasting irradiances. Higher values of total biomass, height and leaf area occurred in sunny than in shaded area. On the other hand, all young plants survived in shade under natural water stress probably by reason of fast and intense biomass accumulation in favor of roots in early development. There was no significant difference about nutrient concentration in leaves between plants growing in sunny and shaded areas. Net photosynthesis in shade increased occasionally when bunches of direct light reached the leaves. Theses sunflecks took place more frequently and at high intensity in dry period but they were more effective for net photosynthesis in rainy period. The ability of young plants to persist under natural conditions in contrasting irradiance up to 1,230 days after sowing could explain the wide distribution of C. langsdorffii in Cerrado physiognomies and in different types of forest.Key words: Cerrado, biomass partitioning, chlorophyll fluorescence, leaf gas exchange, irradiance, sunflecks. resUMoO crescimento e o conteúdo de nutrientes nas folhas foram comparados em plantas jovens envasadas de Copaifera langsdorffii crescendo em área sob irradiância total e em área sombreada sem estresse hídrico. A assimilação de carbono e as relações hídricas foram avaliadas por meio da fotossíntese líquida, da eficiência fotoquímica potencial e do potencial hídrico foliar durante cursos diários no período seco e chuvoso nas duas áreas com irradiâncias contrastantes. Maiores valores de biomassa total, altura e área foliar ocorreram na área sob irradiância plena. As plantas jovens sobreviveram sob estresse hídrico em condições naturais na área sombreada provavelmente em função do rápido e intenso acúmulo de biomassa nas raízes logo no início do desenvolvimento. Não houve diferença significativa em relação à concentração de nutrientes entre as plantas crescendo sob diferentes regimes de iluminação. A fotossíntese líquida sob sombreamento aumentou ocasionalmente quando feixes de luz que atravessaram as copas atingiram diretamente as folhas de C. langsdorffii. Esses feixes de luz ocorreram com maior intensidade e mais freqüentemente no período seco, mas foram mais efetivos para a assimilação líquida no período chuvoso. A habilidade das plantas jovens persistirem em condições naturais sob os dois regimes contrastantes de irradiância até 1.230 dias após a semeadura pode explicar a vasta distribuição de C. langsdorffii nas diferentes fisionomias do Cerrado e em outras formações florestais. palavras-chave: Cerrado, feixes de irradiância, fluorescência da clorofila, partição da biomassa, sobrevivência, trocas gasosas foliares.
ABSTRACT. Enterolobium contortisiliquum occurs in native forests formations from southeast to northeast Brazil, and too in Argentina, Bolivia and Paraguay. The objective of this study was to evaluate shading effects in the early growth of E. contortisiliquum plants. We measured leaf gas exchange, chlorophyll content index, biomass allocation and leaf development in plants in sunny and shaded areas. The experiment was performed in Universidade Federal de Viçosa, campus Florestal, Minas Gerais, Brazil. To determine the effects of different light regimes (full sunlight and shade) on the growth of E. contortisiliquum plants, we divided plants into two groups: plants grown in a greenhouse under full sunlight, and plants grown under 70% shade tissue. E. contortisiliquum plants grown in full sunlight showed an increase in maximum net photosynthesis, root, stem, and leaf dry mass, root length, root/stem ratio, and length of the stem. However, E. contortisiliquum under shade showed larger specific leaf area, leaf area ratio, chlorophyll content, long leaf life span, and slow growth. These ecophysiological adjustments in sunny and shaded areas may explain the wide geographical distribution of E. contortisiliquum in many vegetation types.Keywords: leaf area, leaf development, maximum net photosynthesis.Efeitos do sombreamento sobre o tempo de vida foliar e características funcionais na espécie com ampla distribuição Enterolobium contortisiliquum (Vell.) Morong RESUMO. Enterolobium contortisiliquum ocorre em formações florestais nativas do sudeste ao nordeste do Brasil e também na Argentina, Bolívia e Paraguai. O objetivo deste estudo foi avaliar os efeitos do sombreamento no crescimento inicial de E. contortisiliquum. As trocas gasosas foliares, o índice de conteúdo de clorofila, a alocação de biomassa e o desenvolvimento foliar foram avaliados em plantas sob sol pleno e sombra. O experimento foi realizado na Universidade Federal de Viçosa, campus Florestal, Minas Gerais, Brasil. Para determinar os efeitos de diferentes regimes de luz sobre o crescimento das plantas de E. contortisiliquum, dividimos as plantas em dois grupos: plantas que cresceram em estufa a pleno sol e plantas que cresceram sob sombrite 70%. As plantas de E. contortisiliquum que cresceram em pleno sol apresentaram aumento na fotossíntese líquida máxima, massa seca de raiz, de caule e folhas, comprimento de raiz, razão massa de raiz/caule e comprimento do caule. Entretanto, plantas de E. contortisiliquum sob sombra apresentaram maior área foliar específica, razão de área foliar, conteúdo de clorofila, tempo de vida foliar e crescimento lento. Esses ajustes ecofisiológicos em áreas sob sol e sombra podem explicar a ampla distribuição geográfica de E. contortisiliquum em muitos tipos vegetacionais.Palavras-chave: área foliar, desenvolvimento foliar, fotossíntese líquida máxima.
The structure of the Cerrado may be explained by the competition between woody and herbaceous species. However, the rising CO2 concentration ([CO2]) predicted under current climatic change may modify the ecophysiological responses of woody and herbaceous species owing to functional traits of each group, which may in turn modify vegetation structure as competitive relationships change among species. In this study we examined ecophysiological responses and competition between two cerrado species under elevated [CO2]. We selected an herbaceous alien grass (Melinis minutiflora P. Beauv.) and an endemic woody cerrado species (Hymenaea stigonocarpa Mart. ex Hayne). Hymenaea stigonocarpa individuals were maintained in three plots with different M. minutiflora densities: 0, 50 and 100% in two different [CO2] (380 ppm and 700 ppm) in open-top chambers. Leaf gas exchange, effective quantum efficiency of PSII, chlorophyll content, and growth increased in H. stigonocarpa plants under high [CO2]. The competition with M. minutiflora under elevated [CO2] led to an increase in specific leaf area, leaf area ratio and biomass allocation to shoots in H. stigonocarpa. In contrast, M. minutiflora had a delayed leaf development and high stem dry mass under elevated [CO2]. These changes in growth patterns under elevated [CO2] will modify allocation of resources, improving the competition potential of the woody species over the alien grass species in the Cerrado.
The morphology and the biomass allocation in shoots and leaves were investigated in 15 cerrado tree species with distinct leaf phenology growing under natural conditions. Higher values of leaf/shoot ratio on mass base, individual leaf area, leaf area per shoot, leaf display index, and leaf number per shoot length were found in deciduous than in evergreen species. The differences about shoot-foliage relationship across leaf phenological groups could be explained by plagiotropic shoots on deciduous and by erect shoots in semideciduous and evergreen species. Plagiotropic shoots allow similar irradiance along shoots and high biomass allocation in favor of leaves without foliage self-shading in deciduous tree species. The structural differentiation between short and long shoots was indicated by an exponential relationship between leaf display index and shoot length in all deciduous, in three semideciduous, and in two evergreen species. Therefore, especially in deciduous, the short shoots had higher leaf area per unit of length than the long shoots. The differentiation between short and long shoots depends on the shoot length in deciduous because of the leaf number on shoot is predetermined in buds. Contrastingly, the leaf neo-formation in semideciduous and in evergreen tree species keeps the shoot-leaf relationship per shoot length more constant, because of the foliage being produced according to the shoot growth during the year. In conclusion, the foliage persistence, the shoot inclination, the type of leaf production and the resources allocation between autotrophic and heterotrophic vegetative canopy parts are interdependent in cerrado tree species across different leaf phenological groups.
Variações na morfologia e alocação de biomassa em folhas e ramos foram investigadas em 15 espécies arbóreas de Cerrado com distintas fenologias foliares e crescendo em condições naturais. As espécies decíduas apresentaram maiores valores que as sempreverdes para razão entre massa de folhas e do ramo, área foliar individual, área foliar por ramo, índice de exposição foliar e número de folhas pelo comprimento do ramo. As diferenças acerca das relações ramo-folhagem entre os grupos fenológicos poderia ser explicada pelos ramos plagiotrópicos nas decíduas e pelos ramos eretos nas semidecíduas e sempre verdes. Ramos plagiotrópicos propiciam similar irradiância incidente ao longo do ramo e alta alocação de biomassa em favor das folhas sem auto-sombreamento da folhagem nas decíduas. A variação estrutural entre ramos curtos e longos foi indicada por uma relação exponencial entre o índice de exposição foliar com o comprimento do ramo em todas decíduas, em três semidecíduas e duas sempreverdes. Assim, especialmente nas decíduas, ramos curtos apresentam maior área foliar por comprimento de ramo do que ramos longos. A diferenciação entre ramos curtos e longos depende do comprimento do ramo nas decíduas em função do número de folhas no ramo ser pré-determinado nas gemas. Diferentemente, a neo-formação de folhas nas semidecíduas e semprever...
This study investigated the relationship among shoot inclination, irradiance and morphophysiological traits of basal (BL) and distal (DL) leaves in six cerrado trees with distinct leaf deciduousness. Deciduous species showed plagiotropic shoots (44º) and larger leaf area than semideciduous and evergreen species, which showed orthotropic shoots (56º and 63º, respectively). Despite larger leaf area, irradiance at shoot base in deciduous was around 85% in relation to full irradiance, while in semideciduous and evergreen only 23% of irradiance reached on BL. Likewise, maximum net photosynthesis (P nmax ) was similar between BL and DL in deciduous. Contrastingly, semideciduous and evergreen showed significant decreasing of P nmax in BL. Plagiotropic shoots of deciduous allow similar irradiance along shoots, resulting in similar P nmax along shoot. On the other hand, orthotropic shoots of semideciduous and evergreen resulted in shading and decreasing of P nmax on BL. However, considering BL and DL together, there was not significant difference of leaf-life-span or P nmax among deciduous, semideciduous and evergreen. Therefore, shoot inclination and shading among leaves on same shoot should be significant influencing leaf morphophysiological traits along shoots in cerrado trees. On the other hand, similar leaf-life-span among phenological groups resulted in absence of P nmax differences when BL and DL were considered together.Key words: Leaf-life-span, maximum net photosynthesis, orthotropic, self-shading, plagiotropic. resumoAs relações entre a inclinação do ramo, irradiância e as características morfofisiológicas de folhas basais (FB) e distais (FD) foram investigadas em seis espécies arbóreas de Cerrado com distintas fenologias foliares. Espécies decíduas apresentaram ramos plagiotrópicos (44º) e maior área foliar do que espécies semidecíduas e sempreverdes com ramos ortotrópicos (56º e 63º, respectivamente). Nas espécies decíduas a irradiância solar foi de 85% na base do ramo, mas nas espécies semidecíduas e sempreverdes menos de 23% da irradiância incidiu nas FB. As decíduas apresentaram fotossíntese líquida máxima (P nmax ) equivalente entre FB e FD, porém, nas semidecíduas e sempreverdes ocorreu uma diminuição significativa de P nmax nas FB. Os ramos plagiotrópicos das decíduas permitiram uma equivalente irradiância solar e similar P nmax ao longo do ramo. De forma contrastante, os ramos ortotrópicos das semidecíduas e sempreverdes resultaram em intenso sombreamento e diminuição de P nmax nas FB. Considerando conjuntamente FB e FD, não houve um gradiente significativo dos valores do tempo de vida da folha (TVF) ou P nmax entre decíduas, semidecíduas e sempreverdes. A inclinação do ramo e o sombreamento correspondente entre as folhas são importantes fatores para a determinação de P nmax ao longo dos ramos nas espécies arbóreas de Cerrado. Por outro lado, quando FB e FD foram consideradas conjuntamente não houve diferença de P nmax entre os grupos fenológicos estudados devido à similaridade do TVF.Palavras...
The photosynthetic adjustments of macaw palm (Acrocomia aculeata) were evaluated in 30-day-old seedlings exposed to high and low light environments, and sudden transference from low to high light and comparisons were made with the hardening protocol used in nurseries. Furthermore, we evaluated the responses to long-term exposure (265 days) to high and low light environments. Macaw palm seedlings exhibited an efficient mechanism that maximized light capture under scarce conditions, and dissipated excess energy to avoid damaging to the photosystem II under high light. The seedlings showed low saturation irradiance but no photoinhibition when exposed to excess light. When grown under low light intensities, seedlings presented higher photochemical efficiency and minimized the respiratory costs with positive carbon balance at lower irradiance than hardened seedlings did. The hardening procedure did not appear to be an advantageous method during seedling production. Long-term exposure to either low or high light did not cause significant leaf anatomical adjustments. However, the low light seedlings showed higher leaf area and chlorophyll content than those exposed to higher light intensity did, which enabled shaded seedlings to maximize the captured light. Furthermore, the high non-photochemical dissipation allowed rapid acclimation to excessive light exposure. These responses allow macaw palm cultivation and establishment in very different light environments.
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