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Malvaceae): Micromorphology, Histo-Phytochemistry and Ultrastructure

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Malvaceae): Micromorphology, Histo-Phytochemistry and Ultrastructure THE LEAF SECRETORY APPARATUS OF HIBISCUS SURATTENSIS AND HIBISCUS SABDARIFFA (MALVACEAE): MICROMORPHOLOGY, HISTO-PHYTOCHEMISTRY AND ULTRASTRUCTURE Kashmira Raghu Email address: [email protected] A research report submitted to the College of Agriculture, Engineering and Science, University of KwaZulu-Natal, in partial fulfilment of the requirements for the degree of Master of Science in Biological Sciences. June 2015 Supervisor: Dr. Y. Naidoo Email address: [email protected] Co-supervisor: Prof. A. Nicholas Email address: [email protected] Foliar secretory structures of Hibiscus surattensis and Hibiscus sabdariffa, K. Raghu COLLEGE OF AGRICULTURE, ENGINEERING AND SCIENCE DECLARATION 1 – PLAGIARISM I, Kashmira Raghu, declare that 1. The research reported in this thesis, except where otherwise indicated, is my original work. 2. This thesis has not been submitted for any degree or examination at any other university. 3. These thesis does not contain other person’s data, pictures, graphs, or other information, unless specifically acknowledged as been source from other persons. 4. This thesis does not contain other person’s writing, unless specifically acknowledged as been sourced from other researchers. Where other written resources have been quoted, then: a. their words have been re-written but the general information attributed to them has been referenced: b. where their exact words have been used, then their writing has been placed in italics and inside quotation marks, and referenced. 5. This thesis does not contain text, graphics or tables copied and pasted from the internet, unless specifically acknowledged, and the source being detailed in the thesis and in the references section. Signed …………………. Declaration Plagiarism 22/05/08 FHDR Approved i Foliar secretory structures of Hibiscus surattensis and Hibiscus sabdariffa, K. Raghu ABSTRACT The research presented here forms the basis of the ethnobotanical and ethnopharmacological evaluation of Hibiscus surattensis, which is a widely distributed vegetable and medicinal shrub used by African and Indian traditional practitioners. Using light microscopy together with advanced electron microscopy techniques, the leaf topography and internal structure was examined. A closely related species, Hibiscus sabdariffa was also investigated for foliar and histophytochemical comparisons. Hibiscus sabdariffa is a widely consumed medicinal species with well-known biological activity and known chemical principles. Both species belong to the section Fucaria within the genus Hibiscus. Analyses of foliar secretory tissues showed that both H. surattensis and H. sabdariffa were characterised by capitate trichomes as well as mucilage- producing ducts and idioblasts. Head cells of capitate trichomes were between 5 and 7, while lignified stalk cells occurred in numbers of 2 or 3. A basal cell supporting the trichome was implanted in the epidermis comparative study between the two species showed some variation of chemical composition in trichome head cells. Polysaccharidic, alkaloids, phenolic and acidic lipid components were identified for both H. surattensis and H. sabdariffa whereas H. sabdariffa showed an intense staining of proteinaceous substances. Densities of capitate trichomes varied from emergent to mature developmental stages, with the highest number occurring on the emergent abaxial surface for both H. surattensis and H. sabdariffa. However, a distinct trend was observed for H. surattensis, where a decrease in capitate trichomes with progressive development was associated with the proliferation of mucilage idioblasts, particularly on the mature adaxial surface. Mucilage producing tissues are assumed to be associated with a protective role against dehydration in a number of plant families. Idioblasts observed in H. surattensis were visible leaf surface structures, embedded in the epidermal tissues. They contained considerable amounts of acidic polysaccharides and acidic lipids, and appeared to be implicated in reducing evaporative water loss in fully expanded leaves. Mucilage ducts were identified in vascular tissue within leaf veins. They occurred parallel with the conducting tissue and comprised of an epithelial layer of cells which seemed to be secreting a mucilage into an extracellular lumen. Crystal idioblasts were also identified in tissues of both H. surattensis and H. sabdariffa. The sequestration of calcium oxalate is assumed to be governed and regulated by specialised mesophyllous idioblasts that in turn appeared to be triggered by excess calcium within the plant body to differentiate into crystal forming cells. The supposed function of calcium oxalate crystals includes mechanical support and herbivory avoidance. TEM revealed the cellular processes which involved crystal idioblast development in H. surattensis. This appeared to demonstrate that degeneration of organelles which are assumed to ii Foliar secretory structures of Hibiscus surattensis and Hibiscus sabdariffa, K. Raghu occur when crystals reach maximal proportions. The non-glandular trichome component of each Hibiscus species was diverse in trichome type and might have accounted for differences in leaf texture and the leaf indumentum. The prickly texture of Hibiscus surattensis was attributed to rigid stellate and falcate trichomes as well as restrorse prickles, whereas the leaves of H. sabdariffa were glabrous with few falcate, bi- and trifurcate trichomes found mainly at the base or along the midvein. Preliminary phytochemical experiments which involved methanolic, chloroform and hexane extracts, yielded favourable results, which showed that leaves of H. surattensis and H. sabdariffa were chemically similar, in this regard, the author recommended further investigation into the phytochemical nature of H. surattensis. iii Foliar secretory structures of Hibiscus surattensis and Hibiscus sabdariffa, K. Raghu CONFERENCES CONTRIBUTIONS FROM THIS THESIS: Raghu K., Naidoo Y. and Nicholas A. (2013) Leaf Micromorphological Physiognomy of Hibiscus surattensis and Hibiscus sabdariffa. Microscopy Society of South Africa Proceedings. Vol 43. Pp. 6., Pretoria. ACKNOWLEDGEMENTS: I would like to thank the following people: My supervisors, Dr Yougasphree Naidoo, for the opportunity to participate in her research as well as her mentorship and leadership, and Prof. Ashley Nicholas for his keen botanical wisdom and warm guidance. Members of the Microscopy and Microanalysis Units at Westville and Pietermaritzburg campuses (UKZN), in particular Mr Vishal Baruth for his assistance with electron microscopy techniques. My closest friends, Prelina Munien, Ashlin Munsamy, Preyan Armugam, Shivanee Pillay, who have assisted me with plant collection, data interpretation and have motivated and encouraged me during difficulties encountered. Dr C.T. Sadashiva for assistance with phytochemical aspects of the study as well motivating me throughout. Dr Wynston Woodenberg for assistance with sectioning of fresh leaf material. My parents, brother and family for their unending support and motivation throughout this endeavour. The National Research Foundation for their financial support. iv Foliar secretory structures of Hibiscus surattensis and Hibiscus sabdariffa, K. Raghu TABLE OF CONTENTS Page DECLARATION i ABSTRACT ii CONFERENCES CONTRIBUTIONS FROM THIS THESIS vi ACKNOWLEDGEMENTS vi TABLE OF CONTENTS v LIST OF TABLES viii LIST OF FIGURES ix LIST OF ABBREVIATIONS xii CHAPTER 1: INTRODUCTION 1 1.1 DESCRIPTION OF Hibiscus surattensis L. 3 1.2 DESCRIPTION OF Hibiscus sabdariffa L. 7 1.3 RATIONALE OF THE STUDY 9 CHAPTER 2: LITERATURE REVIEW 11 2.1 PHYTOCHEMISTRY AND MICROMORPHOLOGY OF HIBISCUS 11 2.2 TRICHOMES IN PLANT DEFENCE 13 2.2.1 NON-GLANDULAR TRICHOMES 14 2.2.2 GLANDULAR TRICHOMES 14 2.2.3 CAPITATE TRICHOMES 14 2.3 INTERNAL PLANT SECRETORY TISSUES 16 2.3.1 MUCILAGE IDIOBLASTS AND DUCTS 16 2.3.2 CRYSTAL IDIOBLASTS 19 2.4 TAXONOMY 19 CHAPTER 3: MATERIALS AND METHODS 22 3.1 PLANT MATERIALS 22 3.2 STEREOMICROSCOPY 22 3.3 SCANNING ELECTRON MICROSCOPY (SEM) 23 3.4 TRANSMISSION ELECTRON MICROSCOPY (TEM) 23 3.5 TRICHOME AND MUCILAGE CELL DENSITY COUNTS AND 24 STATISTICAL ANALYSES 3.6 HISTOCHEMISTRY 24 v Foliar secretory structures of Hibiscus surattensis and Hibiscus sabdariffa, K. Raghu 3.7 FLUORESCENCE MICROSCOPY 26 3.8 PREPARATION OF LEAF EXTRACTS FOR PHYTOCHEMICAL 26 TESTS CHAPTER 4: RESULTS 29 4.1 STEREOMICROSCOPY 29 4.2 GLANDULAR CAPITATE TRICHOMES 32 4.2.1 HISTOCHEMICAL OBSERVATIONS OF GLANDULAR 34 CAPITATE TRICHOMES 4.2.2 MICROMORPHOLOGICAL OBSERVATIONS OF CAPITATE 37 TRICHOME DEVELOPMENT 4.2.3 ULTRASTRUCTURE OF GLANDULAR CAPITATE 40 TRICHOMES 4.3 MUCILAGE IDIOBLASTS AND DUCTS 42 4.3.1 HISTOCHEMICAL OBSERVATIONS OF MUCILAGE 45 IDIOBLASTS 4.3.2 ULTRASTRUCTURE OF MUCILAGE IDIOBLASTS AND 47 MUCILAGE DUCTS 4.4 CRYSTAL IDIOBLASTS 48 4.5 NON-GLANDULAR TRICHOMES 50 4.6 PRELIMINARY PHYTOCHEMISTRY 52 CHAPTER 5: DISCUSSION 54 5.1 TAXONOMIC SIGNIFICANCE 54 5.2 LEAF INDUMENTUM 55 5.3 GLANDULAR TRICHOMES 55 5.3.1 CAPITATE TRICHOME MICROMORPHOLOGY 56 5.3.2 DIFFERENCES IN DENSITY OF FOLIAR CAPITATE 56 TRICHOMES 5.3.3 CAPITATE TRICHOME HISTOCHEMISTRY 57 5.3.4 GLANDULAR TRICHOME ULTRASTRUCTURE 58 5.4 MUCILAGE CELLS AND DUCTS 60 5.4.1 MUCILAGE IDIOBLAST HISTOCHEMISTRY 60 5.4.2 DENSITY OF MUCILAGE CELLS ON LEAVES OF H. 61 surattensis 5.4.3 MUCILAGE DUCT ULTRASTRUCTURE 61 5.4.4 FUNCTION OF MUCILAGE IN HIGHER PLANTS 61 5.5 CRYSTAL IDIOBLASTS 62 vi Foliar secretory structures of Hibiscus surattensis and Hibiscus sabdariffa, K. Raghu 5.6 PHYTOCHEMICAL FINDINGS
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