Makixo in the Journal of the Tokyo Botanical Society. In this article he regarded the plant as representing a new family, the Mi tr as temon ace se, distinguish- able from the Rafflesiacese by its having a garaophy lions cylindrical and truncate perianth, a mitriformed stamina!
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A little previously, i. In my then description of this new parasite, I was merely following Making, and did not in detail go into the systematic posi- tion of the plant. Later on, I became strongly of opinion tliat the plant belonged to the Raffle si ace se, and accordingly transferred it to that family Hayata II. As I have said above, Mitrastemon was first referred to the Rarrlesiacete, then was considered to represent a new family, and even to be comparable to the Nepenthacere Making II. Consequently, the systematic position of the plant has been much confused. So far as I am aware, a discussion of this kind on Mitrastenncm lias never been undertaken.
Moreover, there is no work on the anatoruy of the intramatrical tissue, the flower-peduncle, or the structure of the ovules, which collectively make up a group of cliaracters which are as important as any other floral characters for the determination of a systematic position. In this paper, I shall first consider the morphology of the plant both external and internal, especially so far as is necessary for the present descussion and then in its bearings on the systematic position of the plant itself.
External Morphology of MitraMemon, especially with regard to its systematic Position. Roots over 1 cm. The volva is always in colony, separate, or confluent below, short, globose, erect, senile, dark fulvous. Peduncles simple, thick, straight or sometimes slightly curved, terete or subcompressed- ly terete, smooth on surface, provided with scales. Scales opposite in 5 or 6 pail's, imbricate, decussate, concave, margin adnate to the peduncles at the base, forming tubes in the basal portions, where honey -like fluids are secret- ed in abundance.
Flowers are always hermaphrodite and usually regular, but sometimes more or less diclinous, especially in the ealier stage of flower-development. Of the four primordial leaves which compose the perianth, the front and back ones are usually broader than the lateral pair. Makino to the TokyS Bot Mag. Kaioasa- sakii Hay. They are solitary, terminal, subterete. The perianth-tube is hypogynous and firmly attached to the ovary. The perianth is 4-merous, as is seen in the very early stage of the flower-development, and also in fully developed flowers which have some- times the perianth 4-lobed.
The existence of this 4-lobed perianth in Mitras- temon escaped the attention of the previous authors who recorded it as having an entirely truncate tube without any lobe, which lobeless perianth they re- garded as one of the peculiar characters separating the plant from the Raffle- siacese Hayata I. Corolla none. Stamens deciduous, hypogynous, erect, connate into a cylindrical cap, entirely covering the gynseomm, as in Pilostylis aetJiiqpica Lotsy p.
Anthers nume- rous connate into a broad belt-like cap which covers the stigma. Anther- cells, quite irregularly and closely arranged in many series on the above mentioned belt-like cap. They are separated by thin walls, and outer walls are especially very simple. A fibrous layer lacks entirely. They are very like those of Pilostylis Lotsy p. Connective-tip- connate into a cap with a very small opening at the apex.
Ovary, entirely superior, sessile, ovoid, one-celled, with parietal placentas, in numbers, or more in KawasasaJcii Hay.
The ovary is much like that of Apodan- thea? Solms-Laubach I. The placentas are lamelliformed, thick, fleshy. Style, crowning the top of the ovary and articulated to it, erect, short, very thick, conical. Stigma, semi-spherico-conical, umbonate at the top with an obscure groove on one side. In shape, the style and stigma are much like those the female flowers of Pilostylis and Apodanthes Solms-Laubach I.
Ovules, minute, obovoid-ellipsoid, with a short or long funicle, anatropous ; integu- ment, composed of a single layer of cells, as is also the case with the nucellus. The structure of ovules are very much like to that of the Rafrlesia- ce? In the case of a plant which was sent from its native locality, and kept growing in a pot in a green-house here in Tokyo, I have observed that the style which at the base is jointed to the ovary is, when the fruit is fully ripe, separated by a slit along the articula- tion. Seeds minute and very numerous, with a hard reticulated testa like those of rafflesiaceous plants.
Inner Morphology of MUrastemon, especially with regards to its systematic Position. Anatomy of the intramatrical Tissue. The intramatrical tissue of the parasite in the host root is entirely of the kind generally called a " thallus " wluch, in its main parts, exists in the bast of the host root.
It is a formless mass just under the peduncle of a flower in the bast and soon separates into a number of threads running right and left along the long axis of the host root ; anastomosing with each other, and forming a network which completely surrounds the host- mot in the middle layer of the bast and on the outer side of the cambium layer. Flower peduncles are generally found most profusely on roots 1 cm.
From each thread of the network-thallus a number of smaller threads much finer than the preceding are sent out perpendicular to it towards the center of the xyleni. For convenience sake, I shall call a thallus-thread running horizontally along the long axis of the root in the bast-region a horizontal thread, and one running vertically towards the center of the xylem a vertical thread.
The horizontal threads in the bast are com- posed, as is seen in a cross section of the host root, of rounded plasmatic cells in the periphery, but of a elongated ones towards the center, and in most cases there are found near the center some tracheids with spiral markings. The peripheral plasmatic cells, it seems to me, are comparable to the phlom of highly organized plants in their function, wlrile the cental somewhat elongated tracheidal cells resemble the xvlem of the same in their function. Accordingly in every cross section of a region in or near the place where a flower- peduncle stands, we shall invariably find in the bast a number of rounded points, each distinct and remote, or sometimes coming into contact and uniting, every one of which is a cross section of a horizontal thread, arranged in a ring or two all round the xyleru.
The surface view of the network of horizontal threads in the bast is clearly and easily observable with the naked eye in the process of removing the outer parts of the secondary bast. Therefore, the intrarnatrical tissue in the bast forms a cylinder-like body made of reticulated threads, which completely surrounds the xylem outside the cambium layer. It is justly comparable in some respects to that of Cytimis which lies in the bast as a true, perfect, but unreticulated cylinder inside the cambium layer. Lotsy p. Such a structure of the intrarnatrical tissue is considered, at the present time, to be a character peculiar to the Raffle si ace se and never observed in any other family.
The only connec- tion of this tissue to one of another kind which we call a " senker," is to be found in Ar. Accordingly in the cross section of a vertical thread, which is obtainable in a tangential section of the host-root, it comes into view as a lentiform group of cells always laying side by side with a medullary ray, or sometimes laying in a group of bast cells. It is broader and thicker towards the base i. The thread which lies deep within the xylem, after having finished its physiological function, is found dead and buried deep in the xylem. The vertical threads are com- posed only of rounded parenchymatous cells which are rounded at the basal portions but by and by become elongate towards the apex, especially in the apical portions which are composed of a single row of cells.
The horizontal threads become more and more slender as they recede from the flower-region, and show no more differentiation of peripheral paren- chymatous and central tracheidal tissue, and in much remoter regions, say 20 or 30 cm.
This shows how widely they are spread over the host root. They are by no means straight, but always anastomosing and running in a zig- zag manner. In longitudinal sections of the host-root, they appear, not as straight hues, but as interrupted or dotted lines. The vertical threads are usually larger in parts near the floral polster, but become shorter in remoter regions. As the horizontal threads reticulate themselves, so do the vertical threads in the region of the xylem, mostly at their apical portions towards the center of the pith.
There the vertical threads push out formless branches towards the apex of the host-root, along its long axis, parallel to the horizontal threads in the bast. Here we have another kind of horizontal threads which connect the vertical threads in the xylem, making an irregular network. This network is composed of irregular threads composed of single rows of rounded cells which are very much larger than those in other parts of the intramatrical tissue, and are of very irregular amoeboid shape. The vertical threads which run usually side by side with the medullary rays are easily distinguished from their companions by their having elongated rounded cells full of plasmas, with much larger unclei, and smooth and much thinner walls.
The cells of the medullary rays, in contrast to those of the vertical threads, are usually rectangular, and have pitted and tliicker walls. Delafield's Hematoxylin solution. While the horizontal threads in the bast sends inwards, in cross section of the host-root, one vertical thread towards the center, they rjush outwards two or tlnee short branches towards the bark ; but they seem to have no important function, not even of absorption.
In cross sections of a remoter part towards the host-apex, i. The cells, the outer-most one especially, of the thallus of this region are, in almost all cases, com- paratively large, and each, the outermost one especially, contains or sometimes as many as eight nuclei. The nuclei are at one time round and at another dumb-bell- shaped, sometimes curved, fusiform or elongated, and the nuclei in the same cell are of very different sizes, some being small, others quite large. They are, in my opinion, as considered from their being very variable in shape and size, and from the position of their cells, possibly formed, not by indirect, but by direct division.
Here, in the present case, the nuclei take, it seems to me, at first a very irregular amoeboid shape, then curve and fold, at last contract at the middle, and divide into two.
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I have also observed a case where two nuclei thus formed seemed to liave come into contact and probably to liave united again. Whether nuclei thus divided by amitose division have the ability to divide mitosely or whether they liave that ability then first when their union is accomplished, I am not in a position to decide, nor does that question come within the purpose of the present paper.
It is quite probable, how- ever, that this amitose division has been caused by the supply of two much nourishment, just as is seen in the cells of endosperrna and those of a root- bulbulus infected with a kind of mycorrhiza Stkasburger 1. We also find here that the swelling of the bast-tissue is especially large in the floral polster. This is possibly ascribable to the same cause as that to which we ascribe the similar effect in the case of galls Kerner p. The intramatrieal tissue, when fully grown, produces endogeneously a flower-bud, which breaks up the bark of the host-root, and protrudes up- wards a peduncle, just as in the case of a plant of the Kafflesiacese.
Solms- Laubach I. The peduncle of Mitrastemon, as is usually the case with a parasite, is extremely primitive in its anatomical structure. The tissue is formed by nearly round yxirenchyinatous cells which are largest towards the center of the tissue mass and become gradually smaller towards the epiderrn. Inter- cellular spaces are greater towards the center and smaller towards the periphery, and there are practically none at all outside of the ring of vascu- lar bundles.
New evidence from embryology in angio-sperm classification. Cronquist, A. See general references. Dell, B. Notes 5: 71— Anatomy of Pilostyles hamiltonii C. Gardner Rafflesiacae in stems of Daviesia. Emmons, L.
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