Botanica Pacifica. A journal of plant science and conservation 2020.9(2):103-112
Article first published online: 16 OCT 2020 | DOI: 10.17581/bp.2020.09214
Stem and leaf structure of Searsia erosa (Thunb.) Moffett (Anacardiaceae) with systematic, ecological and ethnobotanical implications
Nhlamulo N. Mashimbye 1, Annah N. Moteetee 1 & Alexei A. Oskolski 1,2
1 Department of Botany and Biotechnology, University of Johannesburg, South Africa
2 Komarov Botanical Institute RAS, Saint Petersburg, Russia
The anatomy of stems and leaves of the southern African evergreen shrub Searsia erosa (Anacardiaceae) was studied. This species shows the suite of typical traits of Anacardiaceae, such as the presence of secretory canals in the cortex, secondary phloem, wood rays and vascular bundle of leaf midrib, pericyclic fibres in nearly continuous bands, compound sieve plates on oblique walls, simple perforation plates, alternate intervessel pitting, and septate libriform fibres. Like other Searsia species, S. erosa has abundant sclereids in non-conducting secondary phloem, multicellular peltate glandular trichomes on leaf epidermis, short vessel elements and minute intervessel pits; two latter characters are thought to be diagnostic for this genus. Unlike Asian species of Searsia, S. erosa lacks marginal axial parenchyma as well as prismatic crystals in axial parenchyma and in libriform fibres, but it shows the helical thickenings on vessel walls. The presence of the last trait in a southern African species agrees with association of helical thickenings with the regions that experience water stress. The tangential expansion of secondary phloem in S. erosa and probably in other Searsia species is mainly performed by considerable increase in volume of its cells by their sclerification. Such a way of bark growth in girth is out of scope of bark anatomists. Searsia erosa is distinctive from other congeneric species in its hypostomatous leaves with abundant glandular trichomes on adaxial side. This condition demonstrates a labor division between the adaxial side with glandular trichomes that may contribute to leaf protection, and the stomata-bearing abaxial side providing the gas exchange. Unlike most members of Anacardiaceae, the secretory canals of S. erosa produce the oleoresin containing terpenoids (essential oils) and lipids, but lack polysaccharides. The abundance of glandular trichomes and secretory canals producing terpenoids is a presumable reason of the use of S. erosa in traditional medicine by the Basotho people.
Машимбайе Н.Н., Мотити А., Оскольский А.А. Анатомия стебля и листа Searsia erosa (Thunb) Moffett (Anacardiaceae) в связи с систематикой, экологией и традиционным использованием этого вида. Изучено анатомическое строение стебля и листа южноафриканского вечнозелёного кустарника Searsia erosa (Anacardiaceae). Для этого вида характерен набор признаков, типичных для других представителей семейства Anacardiaceae, таких как наличие секреторных каналов в кортексе, вторичной флоэме, лучах в древесине и в сосудистом пучке главной жилки листа, перициклические волокна в почти непрерывных полосах, сложные ситовидные пластинки на скошенных стенках, простые перфорационные пластинки, очередная межсосудистая поровость, септированные древесинные волокна. Подобно другим видам Searsia, S. erosa характеризуется обильной склерификацией непроводящей вторичной флоэмы, наличием многоклеточных пельтатных железистых трихом на эпидермисе листа, короткими члениками сосудов и мелкими межсосудистыми порами. Вероятно, два последних признака имеют диагностическое значение для рода. В отличие от азиатских видов Searsia, у S. erosa отсутствует маргинальная осевая паренхима, а также призматические кристаллы в клетках осевой паренхимы и в древесинных волокнах, однако имеются спиральные утолщения на стенках сосудов. Наличие последнего признака у южноафриканского вида согласуется с общей приуроченностью спиральных утолщений к регионам, испытывающим водный стресс. Тангентальное расширение луба у S. erosa и, по-видимому, у других видов Searsia, происходит главным образом за счет увеличения объема склерифицированных клеток. Этот способ увеличения периметра коры не привлекает достаточного внимания анатомов. S. erosa отличается от других изученных видов Searsia гипостоматными листьями с многочисленными железистыми трихомами на адаксиальной стороне. Такое строение связано с разделением функций между двумя сторонами листа: верхняя сторона, покрытая трихомами, обеспечивает защиту листа, в то время как нижняя сторона специализирована на газообмене. В отличие от большинства представителей Anacardiaceae, в секреторных каналах S. erosa образуются маслосмолы, в составе которых присутствуют терпеноиды (эфирные масла) и липиды, но отсутствуют полисахариды. Обилие железистых трихомов и секреторных каналов, продуцирующих терпеноиды, может быть причиной использования S. erosa в традиционной медицине басуто.
Keywords: bark, wood, secondary phloem, sclereids, dilatation, hypostomaty, glandular trichomes, histochemistry, secretory canals, terpenoids, oleoresin, Basotho traditional medicine, кора, древесина, вторичная флоэма, склереиды, дилатация, гипостомия, железистые трихомы, гистохимия, секреторные каналы, терпеноиды, традиционная медицина басуто
References
Angyalossy, V., M.R. Pace, R.F Evert, C.R. Marcati, A.A. Oskolski, T. Terrazas, E. Kotina, F. Lens, S.C. Mazzoni, G. Angeles, S.R. Machado, A. Crivellaro, K.S. Rao, L. Junikka, N. Nikolaeva & P. Baas 2016. IAWA List of microscopic bark features. IAWA Journal 37(4):517-615. CrossRef
Barkley, F.A. 1942. A key to the genera of the Anacardiaceae. American Midland Naturalist 28:465-474. CrossRef
Bhatt, J.R. & H.Y. Mohan Ram 1992. Development and ultrastructure of primary secretory ducts in the stem of Semecarpus anacardium (Anacardiaceae). IAWA Bulletin 13:173-185. CrossRef
Buwa, L.V. & J. Van Staden 2006. Antibacterial and antifungal activity of traditional medicinal plants used against venereal diseases in South Africa. Journal of Ethnopharmacology 103(1):139-142. CrossRef
Constabel, C.P., K. Yoshida & V. Walker 2014. Diverse ecological roles of plant tannins: Plant defence and beyond. In: Recent advances in polyphenol research (A. Romani, V. Lattanzio & S. Quideau, eds), vol. 4, pp. 115-142, John Wiley & Sons, Chichester, West Sussex, UK. CrossRef
Crivellaro, A., & F.H. Schweingruber 2013. Atlas of wood, bark and pith anatomy of Eastern Mediterranean trees and shrubs: with a special focus on Cyprus. Heidelberg, New York, Dordrecht, London: Springer. CrossRef
David, R. & J.P. Carde 1964. Coloration différentielle des inclusions lipidique et terpéniques des pseudophylles du pin maritime au moyen du réactif Nadi. Comptes rendus de l'Académie des Sciences, Paris 258(4):1338-1340.
De Vasconcelos, A.L., A.L. De Vasconcelos & K.P. Randau 2016. Pharmacognostic characterization of Spondia mombin L. (Anacardiaceae). Pharmacognosy Journal 8(6):513-519. CrossRef
Dong, Z. & P. Baas 1993. Wood anatomy of trees and shrubs from China. V. Anacardiaceae. IAWA Journal 14(1):87-102. CrossRef
Drake, P.L., H.J. de Boer, S.J. Schymanski & E.J. Veneklaas 2019. Two sides to every leaf: water and CO2 transport in hypostomatous and amphistomatous leaves. New Phytologist 222:1179-1187. CrossRef
Ellis, R.P. 1974. Smodingium dermatitis: the intercellular secretory canals of aerial axis and their relationship to toxicity. Bothalia 11(3):259-262. CrossRef
Eremin, V.M. & A.V. Kopanina 2012. Atlas of the bark anatoof trees, shrubs and lianas of Sakhalin and the Kuril Islands. Poligrafika, Brest, 896 pp. (in Russian). [Еремин В.М., Копанина А.В. 2012. Атлас анатомии коры деревьев, кустарников и лиан Сахалина и Курильских островов. Брест: Полиграфика. 896 с.].
Fahn, A. & R.F. Evert 1974. Ultrastructure of secretory ducts of Rhus glabra L. American Journal of Botany 61:1-14. CrossRef
Feder, N. & T.P. O'Brien 1968. Plant micro-techniques: some principles and new methods. American Journal of Botany 55: 123-143. CrossRef
Gardner, R.O. 1975. Vanillin-hydrochloric acid as a histochemical test for tannin. Stain Technology 50(5):315-317. CrossRef
Gupta, G., & M. Agarwal 2008. Wood anatomy of Anacardiaceae from India with special reference to the systematic position of Rhus. IAWA Journal 29(1):79-106. CrossRef
IAWA Committee 1989. IAWA list of microscopic features for hardwood identification. IAWA Bulletin New Series 10: 219-332.
InsideWood. 2004-onwards. Published on the Internet. [accessed 2020 July 12]. http://insidewood.lib.ncsu.edu/ search. Last accessed 20.09.2020.
Johansen, D.A. 1940. Plant microtechnique. McGraw-Hill, New York.
Jordaan, A. & H. Kruger 1992. Leaf surface and anatomy of two xerophytic plants from southern Africa. South African Journal of Botany 58(3):133-138. CrossRef
Jordan, G.J., R.J. Carpenter & T.J. Brodribb 2014. Using fossil leaves as evidence for open vegetation. Palaeogeography, Palaeoclimatology, Palaeoecology 395:168-175. CrossRef
Joel, D.M. & A. Fahn 1980. Ultrastructure of resin ducts of Mangifera indica L. (Anacardiaceae). II. Resin secretion in the primary stem ducts. Annals of Botany 46:779-783. CrossRef
Kromhout, C.P. 1975. 'n Sleutel vir die mikroskopiese uitkenning van die vernaamste inheemse houtsoorte van Suid-Afrika. South African Department of Forestry Bulletin 50:124.
Lacchia, A.P.S. & S.M.C. Guerreiro 2009. Aspectos ultraestruturais dos canais secretores em órgãos vegetativos e reprodutivos de Anacardiaceae. Acta Botanica Brasilica 23(2):376-388. CrossRef
Li, S., T. Tosens, P.C. Harley, Y.F. Jiang, A. Kanagendran, M. Grosberg, K. Jaamets & Ü. Niinemets 2018. Glandular trichomes as a barrier against atmospheric oxidative stress: relationships with ozone uptake, leaf damage, and emission of LOX products across a diverse set of species. Plant, Cell and Environment 41(6):1263−1277. CrossRef
Madani, I. & A. Farouk 2019. Stem and leaf anatomy of Searsia glutinosa subsp abyssinica (synonym: Rhus abyssina) (Anacardiaceae) from Erkwit, Sudan. European Academic Research 6(11):6194-6199.
McKown, A.D., R.D. Guy, L. Quamme, J. Klápště, J. La Mantia, C.P. Constabel, Y.A. El-Kassaby, R.C. Hamelin, M. Zifkin & M.S. Azam 2014. Association genetics, geography and ecophysiology link stomatal patterning in Populus trichocarpa with carbon gain and disease resistance trade-offs. Molecular Ecology 23:5771-5790. CrossRef
Metcalfe, C.R. & L. Chalk 1950. Anatomy of the dicotyledons, vol. 1. Clarendon Press, Oxford.
Moffett, R.O. 1994. Names of the southern African species of Rhus (Anacardiaceae) and their etymology. Bothalia 24(1):67-76. CrossRef
Moffett, R.O. 2007. Name changes in the Old World Rhus and recognition of Searsia species (Anacardiaceae). Bothalia 37(2):165-175. CrossRef
Moteetee, A. & B-E. Van Wyk, B-E. 2011. The medicinal ethnobotany of Lesotho: a review. Bothalia 41(1):209-228. CrossRef
Moteetee, A., R.O. Moffett & L. Seleteng-Kose 2019. A review of the ethnobotany of the Basotho of Lesotho and the Free State Province in South Africa (South Sotho). South African Journal of Botany 122:21-56. CrossRef
Muellner-Riehl, A.N., A. Weeks, J.W. Clayton, S. Buerki, L. Nauheimer, Y.-C. Chiang, S. Cody & S.K. Pell 2016. Molecular phylogenetics and molecular clock dating of Sapindales based on plastid rbcL, atpB and trnL-trnF DNA sequences. Taxon 65(5):1019-1036. CrossRef
Mugomeri, E., P. Chatanga, S. Hlapisi & L. Rahlao 2014. Phytochemical characterisation of selected herbal products in Lesotho. Lesotho Medical Association Journal 12(1):38-46.
Muir, C.D. 2018. Light and growth form interact to shape stomatal ratio among British angiosperms. New Phytologist 218:242-252. CrossRef
Nair, G.M., K. Venkaiah & J.J. Shah 1983. Ultrastructure of gumresin ducts in cashew (Anacardium occidentale). Annals of Botany 51:297-305. CrossRef
Pell, S.K., J.D. Mitchell, A.J. Miller & T.A. Lobova 2011. Anacardiaceae. In: Families and genera of vascular plants X. Flowering plants. Eudicots. Sapindales, Cucurbitales, Myrtaceae (K. Kubitzki, ed.), pp. 7-50, Springer, Heidelberg, Dordrecht, London, New York.
Ramovha, L.I. 1997. The taxonomic significance of bark structure in South African Anacardiaceae. Unpublished MSc dissertation. University of Pretoria. Pretoria.
Retamales, H. & T. Scharaschkin 2014. A staining protocol for identifying secondary compounds in Myrtaceae. Applications in Plant Sciences 2(10):1400063. CrossRef
Roth, I. 1981. Structural patterns of tropical barks. Borntraeger Gebrüder, Berlin, 609 pp.
Sant'Anna-Santo, B. F., M. Thadeo, R.M. Meira & L. Ascensão 2006. Anatomia e histoquímica das estruturas secretoras do caule de Spondias dulcis Forst. F. (Anacardiaceae). (Anacardiaceae). Revista Árvore 30(3):481-489. CrossRef
Shale, T.L., W.A. Stirk & J. Van Staden 1999. Screening of medicinal plants used in Lesotho for anti-bacterial and anti-inflammatory activity. Journal of Ethnopharmacology 67(1):347-354. CrossRef
Silva-Luz, C.L., J.R. da Piran, J.D. Mitchell, D. Daly, N.V. Capelli, D. Demarco, S.K. Pell & G.M. Plunkett 2019. Phylogeny of Schinus L. (Anacardiaceae) with a new infrageneric classification and insights into evolution of spinescence and floral traits. Molecular Phylogenetics and Evolution 133:302-351. CrossRef
Schweingruber, F.H., A. Börner & E-D. Schulze 2011. Atlas of stem anatomy in herbs, shrubs and trees, vol. 1. Springer, Heidelberg, New York, Dordrecht, London. CrossRef
Schweingruber, F.H., P. Steiger & A. Börner 2019. Bark anatomy of trees and shrubs in the temperate Northern Hemisphere. Springer. CrossRef
Vassilyev, A.E. 2000. Quantitative ultrastructural data of secretory duct epithelial cells in Rhus toxicodendron. International Journal of Plant Sciences 161:615-630. CrossRef
Venkaiah, K. 1992. Development, ultrastructure and secretion of gum ducts in Lannea coromandelica (Hout) Merrill (Anacardiaceae). Annals of Botany 69:449-457. CrossRef
Wagner, G.J. 1991. Secreting glandular trichomes: more than just hairs. Plant Physiology 96:675-679. CrossRef
Weeks, A., F. Zapata, S.K. Pell, D.C. Daly, J. Mitchell & P.V.A. Fine 2014. To move or evolve: contrasting patterns of intercontinental connectivity and climatic niche evolution in "Terebinthaceae" (Anacardiaceae and Burseraceae). Frontiers in Genetics 5:409. CrossRef
Wheeler, E.A., P. Baas & S. Rodgers 2007. Variations in dicot wood anatomy: a global analysis based on the InsideWood database. IAWA Journal 28:229-258. CrossRef
Yang, Y., Y. Meng, J. Wen, H. Sun & Z. Nie 2016. Phylogenetic analysis of Searsia (Anacardiaceae) from eastern Asia and its biogeographic disjunction with its African relatives. South African Journal of Botany 106(1):129-136. CrossRef
Yi, T., A.J. Miller & W. Jun 2007. Phylogeny of Rhus (Anacardiaceae) based on sequences of nuclear Nia-i3 intron and chloroplast trnC-trnD. Systematic Botany 32(2):379-391. CrossRef
Zahur, M.S. 1959. Comparative study of secondary phloem of 423 species of woody dicotyledons belonging to 85 families. Memoirs Cornell University Agricultural Experiment Station 358:1-160.
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