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Artocarpus Forst MORPHOLOGICAL AND ANATOMICAL INVESTIGATIONS ON ARTOCARPUS FORST. I. Vegetative Organs 1, 2 BY M. R. SHARMA (School of Plant Morphology, Meerut) Received June 2, 1962 (Communicated by Dr. V. Puri, F.A.SC.) INTRODUCTION Artocarpus Forst. (Artos--Bread; Carpos--fruiO is a unique genus which, on account of its peculiar and edible fruits in certain species, has attracted attention from very remote past. It is essentially a tropical genus with about 50 species (Comer, 1952 and Jarrett, 1959, a). They are distri- buted from India to China. The Indian species are: A. heterophyllus Lamk., A. lakoocha Roxb., A. altilis (Park) Fosb., A. choplasha Roxb. and A. hirsutus Lain. It is said to be the only Angiosperm whose inflorescence pieces are reported from cretaceous reeks of Greenland (Seward, 1941). It belongs to the subfamily Artocarpoide~e, tribe Artocarp~e which is included in Urticace~e (Hooker, 1885; Ridley, 1924 and Comer, 1952) or in Moracea: (Engler and Prantl, 1881; Hutchinson, 1926; Rendle, 1938; Bailey, 1950 and Lawrence, 1951). Jarrett (1959, a, b) has made a revision of Artocarpus and allied genera touching upon some points of morphological interest also. Wood anatomy of the Morace~e and allied families has been studied in detail by Tippo (1938) and Record and Hess (1940, 1943). Tippo is of the opinion that anatomical specialization in general has proceeded from Moroide~e to Artocarpoidem, Cephaloide~e and Cannaboide~e (Cannabinace~e) and that evolution of floral structures is correlated with the evolutionary development of anatomical structures. The wood of many species of Artocarpus is of great economic importance. A. hirsutus yields one of the best types of wood called ' Aini ", A. chaplasha yields ' Chaplash' and other species yield wood used for various purposes. 1 Based on a part of the thesis accepted for the Ph.D. degiee of the Agra Universily. Research contribution No. 42 ftorn the School of Plant Mc~p~clc~. /¢,~1~1Cc]lete, Meerut. 243 244 M.R. SI-IARMA The genus thus has considerable taxonomic, paleobotanical, anatomical, morphological and commercial interest about it, and can bear some more anatomical and morphological work. In the present communication, atten- tion is focused on the morphological and anatomical features of vegetative organs. MATERIAL AND METHODS The material of vegetative parts of A. heterophyllus Lamk. [syn. A. integrifolius auct., non Linn. f., A. integra (Thunb.) Merr.] and A. lakooeha Roxb., was collected and fixed in F.A.A. from trees cultivated in local gardens and also from Calcutta and Saharanpur respectively. A. ehaplasha Roxb. was collected from Calcutta and Mr. Virendra Prakash Gupta fixed some for me from the gardens of F.R.I., Dehra Dun. Some fixed material of A. altilis (Park) Fosb. (syn. A. ineisa Linn.) was very kindly obtained for me by my colleague Dr. Y. S. Murty from Mr. Veerabhadra Rao of Visakha- patnam. The material of five Malayan species [A. heterophyllus Lamk., A. altilis (Park) Fosb., A. rigidus Blume. (syn. A. eehinatus Roxb.), A. integer (syn. A. champedon Spring.) and A. odoratissimus Blanco.] was very kindly procured through the courtesy of Dr. J. Sinclair of Singapore. The material was dehydrated, cleared and embedded in the usual manner and serial microtome sections of 10-14 microns were cut. Crystal-violet- erythrosin and safranin-fast-green combinations gave satisfactory results. OBSERVATIONS Almost all species of Artocarpus are trees of moderate height of about 50 feet but A. chaplasha may reach 100-150 feet. The main trunk (clean bole) which varies from 6-20 feet in length is thick and stout. Its surface may be rough or smooth, with dark brown or dull grey bark splitting irregu- larly. The main trunk and old branches of A. heterophyllus have irregularly distributed warty excrescences of variable shape and size. Stem of young branches is smooth in A. heterophyllus and A. altilis or rough with wiry long hairs in A. integer or densely hairy in A. rigidus, A. chaplasha and A. odoratissimus, etc. The leaves are stipulate, petiolate, simple and arranged (spirally) on young branches with different angles of divergence. In A. heterophyllus, A. integer, A. altilis and A. chaplasha, etc., the alternate phyllotaxy is of 2/5 type with angular divergence of 144°. In A. lakoocha the leaves are alternately arranged on slender and very much elongated branches which have clear dorsiventral symmetry. They are distichous but the angular divergence Morphological and Anatomical Investigations on Artocarpus Forst. 245 (~d; ARTOCARPUSHETEROPHYLLUS ~i.~i ~''~,~j (?~ , . ,.~, ....,-'-,~,--,.... / I~k ~ ~,~ I:NF c~ . "G ~ Lt Ftas. 1-15. Artocatpus heterophyllus.wFig. 1. External features of a branch (young) showing arrangement of leaves (L), stipules (S) and buds. Fig. 2. Glandular bails (G/q) from stem surface. Figs. 3-11. Serial transverse sections through young shoot showing the diver- gonce of vascular traces to different organs. (LT--leaf traces, ST--stil~ular traces, BT brailch traczs, BTr bud traces.) Note the arrangement of successive young orgars, i.e., leaves (L, I_2, Ls, L4), stipules (S) and inflorescence (INF), etc. Fig. 12. Surface view of leaf epidelmis out- side veins showing star-shaped arrangement of cells around the hair base (/qB) which is impreg- nated by crystals of calcium oxalate. Figs. 13-14. Glandular hails (GH) from leaf. Fig. 15. Surface view of loaf epidermis (Lox~er) showing stomata and glandular hairs. 246 M.R. SHARMA between first and second, and third and fourth leaves is 120 ° and that between second and third, and fourth and fifth is 240 ° and so on. Therefore, the leaves on the dorsal side of the branch are separated from one another by half the distance of what separates them on the ventral side of the branch (Fig. 51). In A. heterophyllus, the leaves are evergreen, thick and coriaceous. In young plants they may be lobed but in adult ones the margin is entire. In A. altilis the leaves are evergreen in Malaya but deciduous in monsoon countries and southerly pacific isles with mild winter (Corner, 1952). In A. lakoocha and A. chaplasha the leaves are deciduous, coriaceous or sub-eoriaceous and larger in size than those of A. heterophyllus. In A. altilis they may be 30-90 cm., deeply dissected or incised into several pointed lobes. Surprisingly enough the leaves appear to be dimorphic in A. chaplasha. The juvenile leaves are up to 90 cm. long, lobed or pinnatifid while in adult condition they are almost elliptical in shape, coriaceous in texture and are not more than 25 cm. in length (Bor, 1953). Each leaf has two caducous and amplexicaul stipules which may be large and spathaceous (A. heterophyllus, A. integer and A. altilis) (Figs. 1, 16 and 25) or small (somewhat semi-amplexieaul) enclosing the bud only for a short duration and becoming free lateral soon after (A. lakoocha) (Figs. 51, 55 and 56). They are mostly hairy and pubescent or glabrous (A. heterophyllus). These stipules protect the young vegetative or flowering buds like a bud scale and soon fall off leaving a prominent characteristic, annular cieatrice (scar) on the node (Figs. 1, 16, 25, 32, etc.). The smaller stipules (A. lakoocha) have small line of attachment and correspondingly their cicatrices are small and when mature do not form a ring. Anatomical features: Internode.--The internode of vegetative shoot is cylindrical and oval or circular i,n cross-section. In flowering shoot it becomes flattened and triangular at a node. The epidermal cells are mostly squarish or radially elongated (A. heterophyllus). The cuticle is thin in A. heterophyllus, A. integer and A. altilis but becomes very thick in A. lakoocha, A. chaplasha, A. odoratissimus and A. rigidus. Some of the epidermal cells are full of latex, more so in A. heterophyllus but many are produced into unicellular and multi- cellular glandular hairs (Figs. 2, 17, 28 and 52). In A. heterophyllus both these types of hairs are small, microscopic and deciduous but in other species only the multicellular glandular hairs are microscopic and deciduous and found on young stem only. They develop from a single epidermal cell by transverse division that cuts off a basal cell. The upper cell divides again transversely and vertically twice and forms a glandular club-shaped head of eight or more cells (Fig. 2). These cells may be full of latex and have thick wall. Some- times a stalk subtends a peltate glandular head of many cells found Morphological and Anatomical Investigations on Artocarpus Forst. 247 mostly in leaves (Figs. 13 and 14). The unicellular hairs of other species, i.e., A. chaplasha, A. rigidus, A. lakoocha and A. odoratissimus, etc., are macro- scopic, dense, rough and may be straight or hooked with pointed tips (Fig. 48). In A. integer they are also tough and pointed but are wiry and much elongated. Because of the dense growth of hairs, the epidermis in these species is irregular or very oftert becomes obscure (Figs: 48 and 50). The cell-wall of these hairs is thick and lignified and has very fine excrescences of cutin and silica of different magnitudes. Thus the hairs appear to have some systematic value. In A. rigidus, A. chaplasha and others with dense hairs, there is an earlier origin of cork cambium. The hypodermis is collenchymatous and chlorenchymatous and has a few layers. Some of their cells have latex in them. Cortex and pith regions are made of parenchymatous cells with conspicuous intercellular spaces. The laticifers are abundant in cortex and pith in A. heterophyllus but their frequency diminishes gradually in A. rigidus, A. altilis, A. integer, A. chaplasha, A. lakoocha and A. odoratissimus. Sph~erocrystals are common in the cortex and pith regions of all the species. In A.
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