EMBRYOLOGICAL STUDIES IN THE FAMILY SAXIFRAGACEAE (S.L.). I. DEVELOPMENT OF THE OVULE AND EMBRYO SAC IN SAXIFRAGA FORTUNEI VAR. PARTITA (MAKINO) NAKAI

The development of the ovule, megaspore and megagametophyte in Saxifraga fortunei var. partita (Makino) Nakai was observed. The ovule is anatropous, bitegmic, and crassinucellate. Both integuments originate from the epidermis. The archesporium is considered to be multicellular. The primary sporogenous cell functions as the megaspore mother cell which forms a T-shaped tetrad. The chalazal member of the megaspore tetrad is functional and develops into a Polygonum-type embryo sac. In the pyriform synergids the filiform apparatus is observed, but any hook or indentations could not be recognized. The antipodal cells are detectable until the Helobial endosperm undergoes several nuclear divisions. Secondary multiplication of the nuclei or the cells of the antipodals could not be observed.     

THE FEMALE GAMETOPHYTE OF THREE SPECIES OF GRINDELIA AND OF PRIONOPSIS CILIATA (COMPOSITAE)

In the four species studied, Grindelia stricta ssp. blakei, G. arizonica, G. lanceolata, and Prionopsis ciliata (Haplopappus ciliatus), the female gametophyte develops according to the Polygonum (normal) type from the chalazal megaspore of a row of four. In most cases only two antipodal cells are formed, the micropylar one originally containing two nuclei. The number of nuclei increases in both cells; one or both antipodal cells typically grow laterally into the integument, often extending to near the surface of the ovule. This resembles the condition previously reported in Grindelia squarrosa. Since the four species of Grindelia have similar antipodal outgrowths, such outgrowths may be considered typical for this genus. The fact that Prionopsis resembles Grindelia in regard to outgrowths from the antipodal cells and differs in this respect from typical Haplopappus helps to justify its separation from Haplopappus, and perhaps its suggested merger with Grindelia.     

獐牙菜的胚胎发生

对獐牙菜大孢子发生、雌配子体形成、受精、胚及胚乳发育过程进行了研究。主要结果如下：子房2心皮,1室,4列胚珠,侧膜胎座;薄珠心,单珠被,倒弯生胚珠。大孢子母细胞减数分裂形成4个大孢子直线形排列,合点端的大孢子具功能,胚囊发育为蓼型。3个反足细胞宿存,每个细胞均多核和异常膨大,反足吸器明显,并在胚乳之外形成染色较深的类似“外胚乳”的结构。珠孔受精,受精作用属于有丝分裂前类型。胚乳发育为核型;胚胎发育为茄型。果实成熟时,种子发育至球形胚阶段。反足细胞在龙胆科一些短命植物中的宿存与分裂具有重要的生殖适应与进化意义。     

DEVELOPMENT OF THE OVULE AND FEMALE GAMETOPHYTE IN EUSTACHYS PETRAEA AND E. GLAUCA (POACEAE)

The unilocular pistil in Eustachys contains a single ovule with lateral placentation. In E. petraea and E. glauca, the mature ovule is bitegmic, tenuinucellate, and amphitropous with the endostomic micropyle oriented toward the base of the locule. A single hypodermal archesporial cell enlarges to form the megasporocyte. The chalazal dyad member is larger than the micropylar one, and meiosis II is nonsynchronized. Two-thirds of the tetrads are linear and one-third T-shaped. The chalazal megaspore is functional. Initially ovoid, the two-nucleate female gametophyte becomes curved as it enlarges. The four-nucleate stage becomes wider at its extremities and constricted in the center. Synchronous mitotic divisions establish the eight-nucleate stage with four nuclei at each pole separated by a large central vacuole. In E. petraea, the maturation sequence begins with antipodal differentiation, followed by differentiation of the egg apparatus, migration of the polar nuclei to the center, and division of the antipodals to produce twelve cells. The sequence in E. glauca begins with migration of the polar nuclei followed by differentiation of the antipodals, egg apparatus, and antipodal replication to six cells. The polar nuclei fuse to form a secondary nucleus appressed to the egg cell in E. glauca and separated from it by a vacuole in E. petraea. T-tests for length measurements for various stages of development indicate that the functional megaspore and two-nucleate female gametophyte are significantly larger in E. glauca than in E. petraea. There is no significant difference in gametophyte length at the four-nucleate stage, and at the eight-nucleate stage, length in E. petraea surpasses that in E. glauca. This gap widens significantly at the mature stage. Nuclear volumes are significantly greater in E. glauca than in E. petraea in the functional megaspore and two-nucleate stage, but the volumes are similar at the four-nucleate stage. Consideration of the differences in structural complexity between the sporophyte and gametophyte generations leads to the conclusion that the female gametophytes of these species are more distinctive than are the sporophytes.     

Ricerche Embriologiche su Senecio Vulgaris L. var. Thyrrenus Fiori

Abstract

Embryological researches on SENECIO VULGARIS L. var. THYRRENUS Fiori. — Male gametophyte, development of tapetal cells and female gametophyte have been studied in Senecio vulgaris L. var thyrrenus Fiori.

1) The development of male gametophyte is normal. Divisions of the microspore mother cells are of the simultaneous type. The division of the generative nucleus has never been observed till the pollen grain was in the anther.

2) The tapetal cells follow a very simple development. The nucleus of each cell divides only twice starting at the same time with the meiotic divisions of pollen mother cells but ending much earlier; subsequently, as usually happens with the Asteraceae, the ameboid involution of the tapetum begins. Endomitosis or any other process which leads to a polyploidy not due to nuclear fusion, has never been observed.

3) The female gametophyte is eight nucleate of the normal type (Polygonum). At maturity it shows only three antipodal cells whose nucleus undergoes at first, two or three divisions. Only later these new nuclei, always within the antipodal cell, may fuse in a polyvalent one.     

Embryological Studies on Ginseng (Panax ginseng C.A. Mey.)

Embrylogical studies of ginseng were carried out systematically. The main results were as follows: 1. The anther is tetralocular. Meiosis of the microspore mother cells was accompanied by simultaneous cytokinesis, and resulting in tetrahedral tetrad. The mature pollen grains are three-celled type. 2. The megasporocyte undergoes meiosis and gives rise to a linear or a "T" shaped tetrad of megaspore, the chalazal megaspore was functional. The development of female gametophyte was of the Polygonum type. 3. In the same flower, the corresponding developmental stages of the stamen and pistil are observed; and we also related them with the outer flower mophology. 4. The double fertilization of ginseng referred to the type of the premitotic syngamy. About 7–10 hours after pollination, most of the female and male nuclei begin to fuse with each other. 5. The primary endosperm nucleus begins the first mitotic division within 36 hours after pollenation, the resting period of the zygote is about 15–20 days long. The endosperm belonged to the nuclear type. When there are about three hundreds free nuclei cell walls begain to appear thus forming cells in the endosperm.     

A CYTOCHEMICAL STUDY OF EMBRYO SAC DEVELOPMENT IN STELLARIA MEDIA

Megasporogenesis and embryo sac development in Stellaria media were investigated using cytochemical methods for the demonstration of nucleic acids, proteins, and polysaccharides. RNA concentrations were high in the archesporial cells, low in the megaspore mother cell, and increased again to high concentrations with the formation of the megaspore and 2-, 4-, and early 8-nucleate embryo sac. RNA levels were also high in the egg and primary endosperm nucleus but low in the synergid and antipodal cells. Nucleolar size and vacuolation were indicative of RNA synthetic activity. Protein concentrations were parallel in concentration and distribution to those observed for RNA. Polysaccharides were conspicuously absent from all stages except the synergids and nucellar cells. Feulgen-stained DNA was demonstrable in the antipodal cells, megaspore mother cell, and megaspore cell, but was not visible in the 2-, 4-, or early 8-nucleate embryo sac. Feulgen staining was also absent from the egg and primary endosperm nucleus but was visible in the synergids and antipodals. Histones were difficult to visualize anywhere except in the egg cytoplasm and the nuclei of the antipodals.     

濒危植物马蹄参的大孢子发生和雌配子体发育

为探讨马蹄参(Diplopanax stachyanthus Hand.-Mazz.)濒危机制与雌性生殖发育的关系,采用石蜡切片法观察马蹄参大孢子形成和雌配子体的发育过程。结果表明,马蹄参雌蕊单心皮,子房下位,1室,1枚胚珠。胚珠横生于短片状胎座上,具单珠被,厚珠心。单孢原细胞自珠心1~2层表皮细胞处分化;大孢子四分体为直线形。成熟胚囊中,2个极核在受精前融合为次生核,3个反足细胞不发达,较早退化;二核胚囊时期出现二核分裂不均且较小核退化消失的异常发育现象。因此,马蹄参雌配子体发育过程中出现异常现象是造成其结实率低的主要原因。     

掌叶大黄胚胎学研究

掌叶大黄(Rheum palmatum L.)的花药4室,单或复孢原。药壁发育为单子叶型。腺质绒毡层发育后期出现双核。小孢子四分体为四面体型,胞质分裂为同时型。成熟花粉为3细胞,表面具3条沟。子房1室,单胚珠,直生,两层珠被,由内珠被形成珠孔,厚珠心。单孢原,位于珠心表皮下。直线形或T形大孢子四分体。合点端的大孢子发育为蓼型胚囊。2个极核在受精前合并为次生核。3个反足细胞宿存。胚乳发育为核型,在球形胚末期开始形成细胞。合点端的胚乳核一直不形成细胞,而为游离核的胚乳吸器。在胚乳吸器和其它部位都发现胚乳核融合现象。胚的发育属于紫菀型。胚具小胚柄。成熟胚囊时期出现承珠盘,且存留时间很长,成熟胚期尚存痕迹。     

POSTFERTILIZATION CHANGES IN OVULES OF MONOTROPA UNIFLORA L. (MONOTROPACEAE)

Monotropa uniflora is an achlorophyllous angiosperm consisting of a mycorrhizae-dependent root system that produces floriferous, aerial shoots. Each of the numerous, minute ovules is anatropous, unitegmic, and contains a Polygonum type female gametophyte. Following double fertilization, a lipid-rich, cellular endosperm develops in association with both chalazal and micropylar haustoria. The vacuolate zygote elongates prior to a cytoplasmically unequal division resulting in a small terminal cell subtended by a larger, vacuolate basal cell. The basal cell eventually degenerates, isolating the terminal cell which is completely surrounded by endosperm. The terminal cell undergoes a cytoplasmically equal transverse division resulting in a two-celled embryo embedded in endosperm. In final stages of seed maturation, lipids decrease and reserve proteins increase in the cytoplasm of both the endosperm and embryo. The morphological reduction of the mature embryo may be associated with a specialized mode of nutrition.     

Female reproductive development and pollen tube growth in diploid genotypes of<Emphasis Type="Italic"> Solanum cardiophyllum</Emphasis> Lindl.

We have characterized female gametophyte (megagametophyte) development and the kinetics of pollen tube growth in self-pollinated diploid genotypes (2n=2x=24) of Solanum cardiophyllum Lindl. that show normal seed formation. In this species megasporogenesis and megagametogenesis give rise to a female gametophyte of the Polygonum type composed of two synergids, an egg cell, a binucleated central cell and three antipodals; however, asynchronous abnormalities resembling mechanisms that prevail during the formation of second division restitution gametes were observed. In self-pollinated pistils at least 1–2% of germinating pollen tubes were able to reach the megagametophyte 60–84 hours after pollination (hap). Although the egg cell acquired a zygote-like morphology 60–84 hap, division of the primary endosperm nucleus was only observed 84 hap. The analysis of genetic variability in full-sib progeny confirmed that seeds are derived from sexual reproduction. These observations suggest that diploid genotypes of S. cardiophyllum can serve as an ideal system to genetically investigate true seed formation in a tuber-bearing Solanum species.     

Lo Sviluppo Del Fiore in Bupleurum Dianthifolium Guss. Con Particolare Riguardo Ad Un Corpo Citoplasmatico Fibrillare Nella Megasporogenesi

Summary

Bupleurum dianthifolium Guss., an endemic species of the small island of Marettimo just West of Sicily, is one of the few constantly hermaphroditic Umbelliferae. It is markedly protandrous. Its chromosome number is n = 8.

The flower develops acropetally from the petals to the stamens, and the carpels; the development of the sepals stops very soon.

The microspores individualise simultaneously and the reproductive nucleus divides while the pollen grain is still in the anther; the tapetum is of the secretory type.

The two carpels fuse at their tips and the ovary is divided into two chambers by the carpophore which develops from the bottom central part of the cavity. Two ovules start developing in each chamber; the upper ones cease their development while the lower ones grow into pendulous, anatropous ovules with only one integument. The ovule is tenuinucellate. The archesporium is unicellular. No parietal cell is formed.

A group of cells trophic in function is present at the basis of the embryo sac, and disappears after fertilisation has occurred. The female gametophyte is of the normal type, monosporic, 8-nucleated, with the antipodal cells sunken in the chalazal nutritive apparatus and disappearing before the polar nuclei fuse. The albumen is nuclear.

A fibrillary body has been observed in the cytoplasm of the megaspore mother-cell. This body after the meiosis remains as an exclusive property of the chalazal megaspore, i. e. the spore that will give the embryo sac. This body can still be seen in the binucleate gametophyte, and disappears during the 4-nucleated gametophyte stage.     

Megasporogenesis, female gametophyte development and embryonic development of Ambrosia L. in China

The megasporogenesis, female gametophyte development and embryonic development of Ambrosia artemisiifolia L. and Ambrosia trifida L. of genus Ambrosia L. in China were studied using conventional paraffin section technology and optical microscopy. The results show that both A. artemisiifolia L. and A. trifida L. have a bilobed pistil stigma, two carpels, one chamber, basal placenta, unitegmic, tenuinucellate, anatropous ovule, and well-developed integumentary tapetum. Megaspore mother cells are directly developed from archesporial cells originated from the nucellar cells under the nucellar epidermis and further undergo meiosis to form linear tetrads. The megaspore at the chalazal end develops into a functional megaspore and the other three megaspores are degraded. The development of embryo sac is monosporic type. After three consecutive mitosis, mononucleate embryo sac becomes a mature embryo sac with two synergids and one egg cell at the micropylar end, a central cell at the center and three antipodal cells at the chalazal end. Most antipodal cells are mononucleate or binucleate, only few are trinucleate. The embryonic development process contains four stages: globular embryo, heart-stage embryo, torpedo-stage embryo and mature embryo. The development of endosperm is cellular type.     

喉毛花的胚胎学研究

本文首次系统地记载了喉毛花属的胚胎发育过程,并以此为依据讨论了该属的分类等级和系统位置。喉毛花花药四室;药壁发育属双子叶型;绒毡层单型起源,细胞具单核,属腺质绒毡层;一层中层细胞;花药壁表皮层宿存,纤维状加厚和膨大;药室内壁减缩。小孢子母细胞减数分裂为同时型,四分体的排列为四面体型;成熟花粉为3-细胞型。子房为2心皮、l室,典型的侧膜胎座,胚珠8列,胚珠胎座靠近两心皮腹缝线。薄珠心,单珠被,倒生胚珠。大孢子母细胞减数分裂形成的4个大孢子呈直列式排列,其中合点端的大孢子具功能。胚囊发育为蓼型。极核在受精前融合为次生核。反足细胞宿存、分裂为8～12个,每个细胞均多核和异常膨大,反足细胞形成的吸器明显。异花传粉,珠孔受精。花粉管通过破坏一助细胞进入胚囊。受精作用属于有丝分裂前配子体融合类型。胚乳发育为核型,每核含2～3核仁。胚胎发育为茄型酸浆I变型,成熟种子胚只发育至球形胚阶段。反足细胞在合子分裂之后才开始退化,在胚的发育过程中反足细胞在胚乳层之外形成一层染色深、类似“外胚乳”的结构。比较喉毛花、龙胆属、假龙胆属以及肋柱花属的胚胎学特征表明喉毛花与假龙胆属的亲缘关系最近,在分类等级上作为一个独立的属较为合适,在系统位置上它比假龙胆属更为原始。     

Female gametophyte development and double fertilization in Balsas teosinte, Zea mays subsp. parviglumis (Poaceae)

Over the course of maize evolution, domestication played a major role in the structural transition of the vegetative and reproductive characteristics that distinguish it from its closest wild relative, Zea mays subsp. parviglumis (Balsas teosinte). Little is known, however, about impacts of the domestication process on the cellular features of the female gametophyte and the subsequent reproductive events after fertilization, even though they are essential components of plant sexual reproduction. In this study, we investigated the developmental and cellular features of the Balsas teosinte female gametophyte and early developing seed in order to unravel the key structural and evolutionary transitions of the reproductive process associated with the domestication of the ancestor of maize. Our results show that the female gametophyte of Balsas teosinte is a variation of the Polygonum type with proliferative antipodal cells and is similar to that of maize. The fertilization process of Balsas teosinte also is basically similar to domesticated maize. In contrast to maize, many events associated with the development of the embryo and endosperm appear to be initiated earlier in Balsas teosinte. Our study suggests that the pattern of female gametophyte development with antipodal proliferation is common among species and subspecies of Zea and evolved before maize domestication. In addition, we propose that the relatively longer duration of the free nuclear endosperm phase in maize is correlated with the development of a larger fruit (kernel or caryopsis) and with a bigger endosperm compared with Balsas teosinte.     

Embryology of Clitoria ternatea (Fabaceae)

ABSTRACT

The embryology of Clitoria ternatea was studied. Anthers contain four sporangia. The anther wall comprises an epidermis, an endothecium, a middle layer and a glandular tapetum. Microspore tetrads are tetrahedral and pollen grains are shed at the 2-cell stage. The ovule is campylotropous, bitegmic and crassinucellate. The micropyle is formed by both the integuments. The megaspore tetrad is linear or T-shaped. The chalazal megaspore is functional and embryo sac development follows the monosporic Polygonum type. Endosperm development is of the nuclear type. The chalazal part of the endosperm forms a haustorium. Embryo development follows the Onagrad type.     

ARTEMISIA PATTERSONII: A 14-CHROMOSOME SPECIES OF ALPINE SAGE

Artemisia pattersonii and A. scopulorum are closely related species of alpine sages endemic to the Rocky Mountains. The species are compared morphologically, ecologically and especially karyologically. Artemisia pattersonii has a chromosome number of n = 7, 2n = 14, which is presently unique in the genus. The karyotype of A. pattersonii is contrasted to its nearest putative relative, A. scopulorum (n = 9, 2n = 18), in order to determine if A. pattersonii might be derived from A. scopulorum through aneuploid reduction; however, the cytological evidence is inconclusive. Possibly these two species are not as closely related as believed previous'y and the nearest relatives of each might occur in north-central Asia.     

Ovule and seed development of Lacandonia schismatica (Lacandoniaceae)

Ovule and seed development is described for Lacandonia schismatica, a species whose androecium is surrounded by the gynoecium. The ovule in each carpel is basal, anatropous, tenuinucellate, and bitegmic. The female gametophyte is formed by the micropylar megaspore cell, after a coenocytic stage of the four megaspore nuclei. The mature female gametophyte has the normal complement of seven cells and eight nuclei. We propose a new type of female gametophyte development on the basis of the coenocytic stage of the tetrad, the cellularization of the tetrad, and the survival of the micropylar spore. At seed dispersal time, the embryo has ~10-20 cells. Endosperm development is of the nuclear type. At maturity, endosperm cells show starch and protein inclusions as well as polysaccharides in their thick walls. The seed coat is formed from the outer integument; the inner one disappears. The exotesta contains tannin. The fruit (achene) wall is two-layered. The maturation of the fruits in a flower is synchronous, and they separate from the receptacle for dispersal.