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Leaf Anatomy of Senna Cana (Fabaceae) in a Seasonally Dry Tropical Forest1

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Leaf Anatomy of Senna Cana (Fabaceae) in a Seasonally Dry Tropical Forest1 Universidade Federal Rural do Semi-Árido ISSN 0100-316X (impresso) Pró-Reitoria de Pesquisa e Pós-Graduação ISSN 1983-2125 (online) https://periodicos.ufersa.edu.br/index.php/caatinga http://dx.doi.org/10.1590/1983-21252021v34n116rc LEAF ANATOMY OF SENNA CANA (FABACEAE) IN A SEASONALLY DRY TROPICAL FOREST1 LUCAS DA PENHA XAVIER2, EMILIA CRISTINA PEREIRA DE ARRUDA2* ABSTRACT - In seasonally dry tropical forests such as the Brazilian Caatinga, factors such as soil type and luminosity are known to cause changes in foliar tissues that enable the survival of species in such places. The present study aimed to describe the leaf morphoanatomy of Senna cana and to observe the plasticity in leaf tissues receiving both direct and indirect light radiation. The study was carried out in the Catimbau National Park in Buíque, PE, Brazil. Mature leaf samples were collected from under full sunlight and full shade in three areas with soils of different colors. Histological slides were prepared for visualization of epidermal faces, internal leaf structure, and histochemical tests. Plasticity was noted in the type of trichome that covered the epidermal faces, and in the number of layers of the palisade and spongy parenchymas, even under the indirect effect of radiation. Phenolic compounds and lipids also showed plasticity. These results demonstrate that Senna cana is a species with high plasticity for the luminosity factor. Further, it was noticed that indirect light also promotes anatomical changes and should be considered in future analyses. Keywords: Caatinga. Semi-arid. Sunlight. Trichomes. Xerophytes. MORFOANATOMIA FOLIAR DE SENNA CANA (FABACEAE) EM UMA FLORESTA TROPICAL SAZONALMENTE SECA Em florestas tropicais sazonalmente secas, como a Caatinga brasileira, fatores como tipo de solo e luminosidade são conhecidos por causar alterações nos tecidos foliares que possibilitam a sobrevivência de espécies nesses locais. O presente estudo teve como objetivo descrever a morfoanatomia foliar de Senna cana e observar a plasticidade em tecidos foliares que recebem radiação de luz direta e indireta. O estudo foi realizado no Parque Nacional do Catimbau em Buíque, PE, Brasil. Amostras de folhas maduras foram coletadas sob plena luz do sol e plena sombra em três áreas com solos de cores diferentes. Lâminas histológicas foram preparadas para visualização das faces epidérmicas, estrutura foliar interna e testes histoquímicos. A plasticidade foi notada no tipo de tricoma que recobriu as faces epidérmicas, e no número de camadas dos parênquimas paliçádico e esponjoso, mesmo sob efeito indireto da radiação. Compostos fenólicos e lipídios também apresentaram plasticidade. Esses resultados demonstram que Senna cana é uma espécie com alta plasticidade para o fator de luminosidade. Além disso, percebeu-se que a luz indireta também promove alterações anatômicas e deve ser considerada em análises futuras. Palavras-chave: Caatinga. Semiárido. Luz do sol. Tricomas. Xerófitas. _______________________________ *Corresponding author 1Received for publication in 05/18/2020; accepted in 10/22/2020. Paper extracted from the monography of the first author. 2Departament of Botany, Universidade Federal de Pernambuco, Recife, PE, Brazil; [email protected] – ORCID: 0000-0002 -8130-1811, [email protected] – ORCID: 0000-0002-1422-2486. Rev. Caatinga, Mossoró, v. 34, n. 1, p. 155 – 165, jan. – mar., 2021 155 LEAF ANATOMY OF SENNA CANA (FABACEAE) IN A SEASONALLY DRY TROPICAL FOREST L. P. XAVIER et al. INTRODUCTION directional cell divisions and is not related to the increase in cell number. Therefore, it is necessary to The Brazilian seasonally dry tropical forest, consider exposure to light when performing studies known as the Caatinga, is the only phytogeographic of ecological anatomy on leaves, which are strongly domain of Brazil that falls entirely within the limits influenced by such a variable. of the national territory (QUESADA et al., 2009; The epidermis is the tissue in direct contact SANTOS et al., 2011). The area was approximately with the external environment and candevelop 844,453 km2 (IBGE, 2004), which borders the specialized cells to provide environmental Cerrado, Amazonian, and Atlantic Forest, which adjustments in plants. Regarding the stomata, Fahn favors different ecotones (QUESADA et al., 2009; and Cutler (1992) consideredamphystomy as a SANTOS et al., 2011). Regarding climate, the standard feature for plants in sunny and xeric sites. A Caatinga presents very extreme conditions in possible explanation is the improvement of the contrast to other Brazilian vegetation formations, conductance of gases within the leaf, made possible with the highest insolation, temperature, and by the presence of the stomata on the adaxial (upper) evapotranspiration as well as lower relative humidity side of the leaves (GUERFEL et al., 2009; YOO et and precipitation (LIMA et al., 2012). al., 2010; FAN et al., 2013). Under such conditions, Most of the Caatinga is located in the an elaborate indument of trichomes capable of northeast portion, with a dominance of crystalline enhancing ultraviolet ray reflection and avoiding rocks that were elevated by tectonic events that water loss through sweating (KARABOURNIOTIS occurred during the tertiary period, and that et al., 2020), and a thick cuticle with the same originated the inselbergsthrough erosive processes purpose (YANG et al., 2011) are commonly (LOPES-SILVA et al., 2019). Araújo-Filho et al. exhibited. (2017) reported that Caatinga’s soils are shallow and Plant secondary metabolitesimprove plant rocky outcroppings are frequent and show different growth and survival under different environmental colors related to its chemical and stresses (ASHRAF et al., 2018). Substances such as physicochemicalproperties. Similarly, according to phenolic compounds can reduce the passage of light Prado (2003), pediments covered by rocks, and and the deleterious effects caused by radiation rocky outcroppings called "lajedos" are typical, and (IZAGUIRRE et al., 2007). Barros and Soares latosols originating from the action of climate on the (2013) considered herbivory and photo-protectionas parent rock occur frequently. The geological essential functions for plants during drought. formation process of the Caatinga resulted in varied Different plant taxa can adapt to local environmental and patchily distributed soils (SANTOS et al., 2013), conditions, and semi-arid regions around the world which have influenced the vegetation. Latosols and exhibit high levels of species richness. nearby sandy soils can contain the same plant species Several floristic inventory studies carried out with different-sized individuals (PRADO, 2003). in the Caatinga have shown that the Fabaceae family Schlichting (2002) found that the morphology is the most representative, followed by of plants is genetically determined, but can be Euphorbiaceae (ALCOFORADO-FILHO, 2003; influenced by the environment. In xeric areas, it is ROCHA et al., 2004; RODAL et al., 2008; GOMES; common for plants to develop smaller leaves to limit ALVES, 2010; MACHADO et al., 2012; transpiration and ameliorate the adverse effects of SANTANA et al., 2016). Fabaceae are dominant high temperatures (FAHN; CUTLER, 1992; even in areas with different soils (COSTA et al., FIGUEIREDO et al., 2015). However, they may be 2015) and distinct phyto-physiognomies (ARAÚJO thickened due to the accumulation of photosynthetic et al, 2011), which indicates a high resistance tissues as well as due to storage of water in the capacity to local stresses.Senna cana (Nees& Mart.) mesophyll (BOEGER; GLUZEZAK, 2006). H.S.Irwin & Barneby, analyzed in this study, is a One of the leading causes of stress in plant native, episodic, evergreen, and a potentially species in the Caatinga is the high level of exposure medicinalspecies. Itisfound in Caatinga, Campos to ultraviolet-B radiation, which is capable of Rupestres, Cerrado, and rocky outcroups slowing the growth of individuals and decreasing (MONTEIRO, 2012), andcan be an ideal model leaf area (ZHAO et al., 2020). This damage varies species for ecological and plasticity studies. according to the degree of exposure of the leaves, Although the family has been well studied, with leaves closer to the ground being less little is known about the anatomy of the group, susceptible to the negative effects of ultraviolet-B especially for seasonally dry tropical forests. Further radiation due to shading (PARISI et al., 1998). studies are required to describe and understand not Furthermore, leaves growing in the sun are known to only the tissue structure of representative taxa, but exhibit marked morphological differences when also the physiology and ecology of these species. compared to those in the shade. In general, they are This study aimed to characterize the foliar thicker and possess more layers of palisade morphoanatomy of individuals of Senna cana parenchyma in relation tospongy parenchyma, (Fabaceae) present in the Caatinga, and to identify which, according to Kim et al. (2005), occurs due to plastic anatomical variations related to the color of 156 Rev. Caatinga, Mossoró, v. 34, n. 1, p. 155 – 165, jan. – mar., 2021 LEAF ANATOMY OF SENNA CANA (FABACEAE) IN A SEASONALLY DRY TROPICAL FOREST L. P. XAVIER et al. thesoil in which these plants were established as well sampled from the fourth node of nine adult as the degree of light incident on the leaves. individuals of Senna cana. Leaf samples exposed to sunlight and shade were collected from the same individuals.
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