McCarrny.—Structure and Development of Astelia nervosa 343 The Structure and Development of Astelia nervosa var sylvestris. By Exata M. MoCarruy, M.Se. (Communicated by Dr. J. B. Holloway.) [Read before the Otago Institute 8th May, 1928; received by Editor, 42th May, 1928; rssued separately, 0th August, 1928.] Prare 52, ‘Tue following account of certain aspects of the structure and develop- ment of Astelia nervosa var. sylvestris is taken from a thesis written at the Botany Laboratory, University of Otago, under the direction of Mrs. M. W. Aitken, M.Sc. It owes its present form to the help and encouragement extended by Dr. J. E. Holloway. Astelia nervosa var. sylvesiris is stout perennial herb growing abundantly under mesophytie conditions on low hillsides in both islands. The leaves are arranged in dense tufts, a number of which combine to form the large head which comprises the plant. The* growth-form of the Asteliads is very similar but Astelia nervosa can be distinguished from other species by its orange berry, the perianth of which is enlarged and coloured. ‘The leaf is long and narrow, varying in length from 3 — 5 ft. and in breadth from 2 — 3 inches.’ It is many nerved but three nerves are very conspicuous, and are coloured red with anthocyanin, which is dissolved in the cell-sap of the superficial cells. The leat as a whole is flexible, but the base is white and succulent and elothed by a tomentum of long silky hairs. Astelia nervosa var. sylvestris is very similar to Astelia Cockaynet (Astelia nervosa var. montana). The latter is found mostly in sub- alpine situations as a small sturdy plant covered with hairs. The structure of the leaf of Astelia Cockaynei as found in the Peridotite ‘Belt, Nelson, is described in ‘‘ Notes on the Autecology of Certain Plants of the Peridotite Belt, Nelson’’ (Trans. N.Z. Inst., vol. 52, pp. 305-308) by Miss M. Winifred Betts, M.Sc. It is practically the same as that of Astelia nervosa var. sylvesiris. It also has some resemblance to Astelia Solandri, as described by Miss J. H. Wilson, M.Se. in ‘‘ Some Plants from the Lava-Field at Mt. Wellington” (Trans. N.Z. Inst., vol. 58, pp. 259-263). This resemblance lies chiefly in the anatomy and function of the leaf bases (Wilson, p. 262). The only other paper dealing with the anatomy of the Asteliads is one by Miss E. M. Herriott, M.A. (Trans. N.Z. Inst., vol. 38, pp. 377-422), “‘ On the Leaf Anatomy of Astelia linearis var. sublata.’? This leaf has no close resemblance anatomically to that of Astelia nervosa var. sylvestris. Mioroscorrc Structure or Marure Lear. (Figs. 1-6). Over the midrib the upper epidermis is composed of small cells, ‘deneath which is an aqueous tissue consisting of 4-6 layers of colour.344 Transactions. less palisade-shaped cells very regularly arranged. The chloren- chyma is below this, and is represented by about 10 layers of spongy parenchymatous cells. The lower epidermis consists of 2 rows of small colourless cells, The fibro-vascular bundle forming the midrib, Fic. 2.—Transverse section of midrib X 140.McCarruy.—Structure and Development of Astelia nervosa 345 stretches from the aqueous tissue to the lower epidermis, where it forms a conspicuous ridge on the under-surface of the leaf. The upper flange of mechanical tissue is a crescent-shaped mass of selerenchymatous cells. It is smaller than the lower flange which would be circular but for a small area occupied by the phloem. Fic. 4.—Transverse section of leaf at margin X 200. ‘The most prominent nerves of the leaf are the two lateral ones. This is due to a very conspicuous anchor-shaped girder of mechanical tissue stretching from one epidermis to the other. The upper flange is the larger and the vascular bundle occupies only the lower half of the web. The aqueous tissue is slightly wider near these veins, and the cells are almost spherical and loosely arranged. Two layers346 Transactions, extend to the web of the girder underneath the upper flange. Towards the midrib the chlorenchyma increases greatly in thickness the upper limits being almost parallel to the surface of the leaf Fic, 6.—Multicellular hair from base of leaf X- 125, which here projects as a ridge. Towards the margin of the leaf the epidermal cells become larger and the aqueous cells smaller. The aqueous tissue is here only one layered, the chlorenchyma occupying most of the leaf. Between each bundle is a large colourless cell several times larger than an ordinary cell of the chlorenchyma. ‘The fibro-vascular bundles are similar to all the other minor bundles of the leaf, that is, they have mechanical tissue in the form of two crescent-shaped areas, one above, one below the vascular bundle. A slight variation oceurs in the end-bundle where the upper flange is practically undeveloped. At the base the leaf is soft and white and covered with a tomentum of silky hairs. These occur on both surfaces of the leaf, but are more numerous on the inner or upper surface. The hairs arise below the level of the epidermis and consist, below the epidermis, of a short stalk 2 or 8 cells deep and 2 wide. Beyond the level of the epidermis the hair branches out in several directions, but as a whole continues at right angles to the surface, branches being given off to the right and left. The branches, which are unicellular, remain close to the parent axis. On some hairs a number of branches areMcCarrny.—Structure and Development of Astelia nervosa 347 given off together forming a whorl. The mesophyll is composed of round loosely-arranged cells which near the upper epidermis form a sponge-like tissue with large intercellular spaces. The three main veins are not conspicuous, but above each large vein is a triangular mass of collenchyma. Below the veins are crescent-shaped gum- passages, The gum is apparently secreted by special mucilage vesicles which project into the gum-passages. These passages, unlike those of A. Solandri (Wilson p. 262), are found only in the leai-base. Microscopic Srrucrure or Lear or Seeptanu. (Fig. 7.) The leaf of a 4-inch-high seedling is slightly thicker in the proximity of the main vascular bundles. On the upper surface there are 2 layers of colourless polygonal cells, the lower representing the aqueous tissue, The chlorenchyma which consists of from 3-6 layers of cells is in the form of spongy parenchyma. The lower epidermis consists of a single layer of cells except near the midrib where there are two layers. At this stage the leaf has 7 veins. These are the three main veins with a small group of sclerenchymatous cells above and below the vascular strand, two fairly large veins with no348 Transactions. mechanical tissue, and two small veins between the main laterals and the midrib. At'the base of the leaf there are no gum-passages. These appear together with the hairs when the plant is about 2 ft. high. . Fic, 7.—Transverse section of young leaf X 50. Microscopic STRUCTURE OF PEDUNCLE OF INFLORESCENCE (Fig. 8). ‘The triangular peduncle has a ground-tissue of round cells rich in starch. The outer layers contain a few chloroplasts and are surrounded by a dermis of tabular-shaped cells. A’oout 1/50 - 1/75 “0 6 0 6 S O° d Oyyr ea 3 4 scbovobecolessD 20200» »afocalo0™ Fie. 8—Diagramatic section of peduncle of flowering shoot X 12.McCarrny.—Structure and Development of Astelia nervosa 349 of an inch from the dermis and parallel to it lies a row of vascular strands. Along this row large vascular bundles occur at fairly regular intervals, the space between each being oceupied by 2 or 3 smaller ones. Inside this row vascular bundles are seattered through the ground-tissue. The mechanical tissue, especially in plants grown in the shade, is not well developed. Frowsrine (Figs. 9 and 10). Cheesoman (p. 317) states that Astelia nervosa is a dioecious plant, All the specimens examined by the writer, however, seemed to be hermaphrodite. This was confirmed by microscopic examina- tion, All the anthers contained pollen-grains and in most cases the ovaries contained ovules. In some cases however the ovules were not well developed and were possibly functionless. An examina- tion of the plants in the fruiting season shows a luxuriant develop- ment of fruit on some plants, while others have none at all. This would certainly seem to indicate the dioecious habit. Astelia nervosa var, sylvestris, however, does not flower and fruit every year. Investigation over a number of years is necessary before the position can be stated definitely. Fra. 9.—Individual flower X 6. Fira. 10—Small portion of flowering spike (nat. size). About February and March individuals about to flower appear slightly swollen at the base. The inflorescence is formed at the apex of the shoot, a new shoot arising laterally to carry on the growth of the tuft. The flowering-seape appears above ground at’ the begining of September, and when only 4 or 5 inches long the pollen-grains and ovules are already formed. The buds open shortly after they emerge from the bracts and surrounding leaves, so that tho tip of the spike is often in full bloom before the rest of the spike350 Transactions. has appeared. The flowers are not conspicuous, the colours being very dull, but they have a sweet scent. In contrast to the flower the fruit is very conspicuous, each spikelet being crowded with brilliant orange berries. The berries soon drop off when the succulent perianth, which at first is closely pressed to the berry, turns back. The number of seeds per berry varies from 3-10, 8 being the most frequent number. The seeds are smooth, black, and angular, with a bright polished appearance. The testa is very hard, thick, brittle, and practically impermeable to water. Srrucrure anv Devevormenr or Antuer (Figs. 11 and 12). In a very young flower, when the whole inflorescence is about an inch long, the stamen is quite undifferentiated, being represented by a mass of homogeneous tissue, kidney-shaped in transverse Fie. 11.—Transverse section of young anther X 500. section. A longitudinal section at the same stage shows a cone-like structure with no distinction between anther and filament. A little later some of the cells of the hypodermal layer divide to form the primary wall-layer and the primary sporogenous layer. This cell- division takes place, as in most Angiosperms, in 4 regions of the anther. Transversely the anther is still the same shape, but in longi- tudinal section shows a slight constriction at the base. This is the beginning of the differentiation into anther and filament, The primary sporogenous layer divides until several layers of sporogenous cells are formed. Some of the wall-layers form a tapetum which is partially disorganised in the spore-mother-cell stage. In longitudinal section at this point the anther and the filament are quite distinct.McCarrny.—Structure and Development of Astelia nervosa 351 The filament, however, is very short while the anther is almost the mature size. “The opening of the bud therefore entails a great elonga- tion on the part of the filament. The stamens pass the winter in the spore-mother-cell stage. ‘At the beginning of spring the spore-mother cells divide to give the tetrad of pollen-grains which have sculptured coats. Directly below the epidermis of the anther is-the endothecium which is com- posed of a layer of palisade-cells. Between the adjacent sporogenous tissues of an anther lobe are 2 or 3 layers of thin-walled cells. In dehiscence this thin-walled tissue breaks down with the result that there is now a single compartment in the anther-lobe. In the region Fic, 12.—Transverse section of anther X 100, showing the coalescence of the 2 adjacent pollen-chambers. of the thin-walled cells the epidermis and endothecium are bent inwards, the apex of the bend being occupied by the thin-walled tissue. ‘Consequently when this breaks down, the wall is not con- tinuous, and with a slight increase of pressure may be bent back- wards setting the pollen-grains free. Srrucrore aNp DeveLopment or Ovuxe (Figs. 13-19). The ovule originates in the ovary about the same time as the division to form the spore-mother-cells occurs in the anther. The mucellus appears first as a protuberance on the placenta. A transverse section at this stage shows the three carpellary leaves with the ovules as small outgrowths at their margins. As the ovule develops the nucellar mass protrudes further and gradually bends sideways and downwards. The first integument soon appears as a ring round the base of the nucellus. Later, but before the first integument envelops