DEVELOPMENTAL MORPHOLOGY OF THE EMBRYO AND SEEDLING OF RHIZOPHORA MANGLE L. (RHIZOPHORACEAE)

The embryo of Rhizophora mangle L. is initially attached to the integument by a long multiseriate suspensor. Its basal cells lyse, and intrusive growth of the endosperm envelops the embryo, forces the micropyle open, and often carries the embryo out of the integument. Thus, “germination” is effected by growth of the endosperm rather than of the embryo. The surface of the endosperm differentiates into a layer of peculiar transfer cells. The cotyledonary body initiates as a toroidal primordium, which later becomes lobed; most of the free portions ultimately fuse. After “germination,” the axis of the viviparous seedling grows by a diffuse intercalary meristem below the cotyledonary node. Before seedling abscission, the shoot apex produces three pairs of leaves, the first of which aborts, leaving the rest of the plumule protected by their stipules. The (immersed) radicle apex is nearly inactive, but lateral roots arise early in seedling development; these are usually the first or only roots to grow during establishment. Ten provascular strands “differentiate” in the cotyledons; a hollow provascular cylinder develops in the hypocotyl. Initial vascular differentiation in the latter is of many alternate poles of xylem and phloem; later, de novo differentiation of metaxylem opposite the protophloem poles, and vice versa, produces collateral bundles. Xylem maturation is endarch over most of the length of the hypocotyl, but tangential and random series of metaxylem vessels occur in the radicle end.     

大车前胚乳发育的超微结构研究

采用透射电镜技术对大车前(Plantago major L.)胚乳发育的超微结构进行了研究。结果表明:(1)大车前为细胞型胚乳;初生胚乳核经一次横分裂产生1个珠孔室细胞和1个合点室细胞;珠孔室两次纵向分裂一次横向分裂形成2层8个细胞,位于上层的4个细胞发育为4个珠孔吸器,位于下层的4个细胞发育为胚乳本体;合点室细胞进行一次核分裂,发育为两核的合点吸器。(2)珠孔吸器呈管状插入珠被组织,珠孔端细胞壁加厚呈现少量分支并具有壁内突,壁内突周围细胞质里分布着大量线粒体、粗面内质网、高尔基体、质体等,细胞核与核仁明显,细胞质浓厚,代谢活动旺盛;球胚期,珠孔吸器的体积呈现最大值,珠孔吸器周围的珠被组织均被水解,形成明显的空腔。珠孔吸器从珠被组织吸收并转运营养物质至胚乳本体,参与胚乳的构建与营养物质的贮藏。球胚后期,珠孔吸器逐渐退化。(3)4个胚乳本体原始细胞具旺盛的分生能力,经不断的平周与垂周分裂增加胚乳细胞数目,使胚乳本体呈现圆球体状,并将胚包围其中;珠孔吸器、合点吸器以及珠被绒毡层吸收转运的营养物质贮存在胚乳本体;球胚后期,随着胚柄的退化,胚体周围的胚乳细胞被水解,为发育的胚所利用。(4)合点吸器的2个细胞核与核仁巨大,线粒体、质体、高尔基体、内质网主要绕核分布,液泡化明显;胚体与胚乳本体的体积增大,逐渐将合点吸器向胚珠合点部位挤压,合点吸器周围的合点组织逐渐被水解,形成巨大空腔。合点吸器自珠心组织吸收并转运营养物质至胚乳本体,参与胚乳的结构构建与营养物质的贮藏。球胚后期,合点吸器逐渐失去功能,呈现退化状态。     

Transfer Cells in Suspensur and Endosperm during Early Embryogeny of Vigna sinensis

During early embryogeny, structural differentiation of the suspensor and endosperm can be observed with the formation of cells with wall ingrowths. In the early proembryo stage, wall ingrowths are seen only on the boundary walls of the embryo sac around the proembryo and at the chalazal end. Later, ingrowths appear in the outer walls of the basal suspensor cells and some wall ingrowths also begin to develop in the outer walls of cellular endospermic cells adjacent to the nucellar cap and the inner integumentary tissues. The suspensor appears to remain active throughout the differentiation stages. Two regions can be clearly distinguished in the suspensor: a basal region and a neck region. Wall ingrowths appear to form only in the cells of the basal region. During the development of the cellular endospermic sheath, its cell number and size both increase slightly. Later, these cells rapidly become separated from each other. Those endospermic cells that abut directly onto the integumentary tissues also develop wall ingrowths. In the region of the fluid endosperm, wall ingrowths are especially abundant in the boundary walls on the ventral side of the embryo sac. The possible pathway of nutrient flow to the developing embryo is discussed.     

花生胚乳细胞化的超微结构观察

花生（ＡｒａｃｈｉｓｈｙｐｏｇｅａｅＬ．）心形胚期的胚乳游离核多瓣裂,或具长尾状结构。胚乳细胞质内有大量线粒体、质体、高尔基体、小泡及少量内质网。中央细胞壁有壁内突。球胚及心形胚期常见胚乳瘤。心形胚晚期,胚乳开始细胞化,胚乳细胞壁形成有３种方式,分别存在于不同的胚珠中：（１）从胚囊壁产生自由生长壁形成初始垂周壁,具有明显的电子密度深的中层,其生长主要靠末端的高尔基体小泡及内质网囊泡的融合。两相邻的自由生长壁末端或其分枝末端相连形成胚乳细胞。（２）核有丝分裂后产生细胞板,细胞板向外扩展并可分枝。间期的非姊妹核间也观察到形成了细胞板。小泡与微管参与细胞板的扩展,高尔基体和内质网是小泡的主要来源。细胞板的扩展末端相互连接,形成胚乳细胞的前身。小泡继续加入细胞板的组成,以后形成胚乳细胞壁。（３）胚乳细胞质中,出现一些比较大的不规则形的片段性泡状结构,它们可能来源于高尔基体小泡,这些片段性泡状结构随机相连形成细胞壁,未见微管参与。胚乳细胞外切向壁及经向壁上有壁内突。     

Ultrastructural characteristics of « Diplotaxis erucoides (L.) DC. » suspensor

Abstract

The development and general morphology of Diplotaxis erucoides (L.) DC. suspensor is of the « Onagrad Type », « Alyssum Variation ». Maximum growth of the suspensor occurs from the globular to the early heart stage of embryo development. The suspensor starts then to degenerate disintegrating shortly after the torpedo stage of the embryo.

The wall ingrowths of the long, tapering, basal cell are especially abundant at the cell's micropilar pole which is closely surrounded by well developed wall ingrowths formed by the endosperm. Wall ingrowths and plasmodesmata are present on the suspensor cells cross walls with the exception of the cell closest to the embryo. No such structures in fact are present on the walls separating this last cell both from the embryo and from the rest of the suspensor. Wall ingrowths are generally associated with numerous, large, mitochondria.

The morphological data seem to indicate that absorption and transport of nutrients from the surrounding tissues is a main function of the suspensor. The possibility of an elaborative and secretory function of this structure is discussed.     

白刺胚乳早期发育的超微结构研究

白刺(Nitraria sibirica)胚乳发育经历游离核阶段、细胞化阶段和被吸收解体阶段。游离核胚乳沿胚囊壁均匀排列为一层,胞质浓厚,其中有丰富的质体、线粒体、高尔基体、内质网和各种小泡等细胞器。珠孔区域的胚囊壁具发达的分枝状壁内突,而周缘区域的胚囊壁具间隔的钉状内突,内突周围的细胞质中具多数线粒体和小泡。胚乳细胞化时,初始垂周壁源于核有丝分裂产生的细胞板。在细胞板两端开始壁的游离生长,一端与胚囊壁相连接,另一端向心自由延伸。壁的游离生长依赖于小泡的融合。早期胚乳细胞具大液泡,具核或无核,细胞质中有大量的线粒体,质体缺乏,其壁仍由多层膜结构组成。     

Functional Anatomy of the Ovule in Broad Bean (Vicia faba L.): Ultrastructural Seed Development and Nutrient Pathways

Ovules of broad bean (Vicia faba L.) were studied to discloseultrastructural features, which can facilitate nutrient transportto the embryo sac from 10 d after pollination (DAP) to the matureseed. Fertilization occurs during the first 24 h after pollination.The endosperm is a coenocyte, which is eventually consumed bythe embryo. By 10 DAP the inner integument is degraded and theouter integument adjoins the embryo sac boundary. The heart-shapedembryo approaches the embryo sac boundary at two sites, whichhere are named contact zones. Small integument cells in theneighbourhood of the first formed contact zones become separatedby prominent intercellular spaces. A heterogenous scatteringmaterial, probably representing secretion products accumulatesin these spaces. By 14-16 DAP the integument exudate disappears,and the suspensor degenerates. As the contact zones increasein size, wall ingrowths form a bridging network in the narrowspace between the embryo sac boundary and the extra-embryonicpart of the endosperm wall. The epidermal cells of the embryoseparate adjacent to these zones, and develop conspicuous wallingrowths. At 20 DAP vacuoles showing various stages in formationof protein bodies appear in the cells of the embryo.Copyright1994, 1999 Academic Press Vicia faba, broad beans, ovule, seed, nutrient transport     

Embryo and Endosperm Development in Isatis Tinctoria L. and the Histochemical Study of Its Storage Reserve

The ovule is anatropous and bitegmic. The nuceIlar cells have disorganized except the chalazal proliferating tissue. The curved embryo sac comprises an egg apparatus and a central cell with two palar nuclei and wall ingrowths on its micropylar lateral wall. The antipodal cells disappear. Embryo development is of the Onagrad type. The filament suspensor grows to a length of 785 μm and degenerats at tarpedo embryo stage. The basal cell produces wall ingrowths on the micropylar end wall and lateral wall. The cells of mature embryo contain many globular protein bodies, 2.5–7.5 μm in diameter, composed of high concentration of protein and phytin, insoluble polysaccharide and lipid. The cells, except procambium, also contain many small starch grains. Some secretory cavities scattered in the ground tissue have liquidlike granules composed of protein, ploysacchaide and lipid. Endosperm development follows the nuclear pattern. At the late heart embryo stage, the endosperm around the embryo and the upper suspensor and the peripheral endosperm of the basal region of the U-shaped embryo sac becomes cellular. The endosperm at micropylar and chalazal ends remains free nuclear phase until the late bended cotyledon stage. Wall ingrowths at both micropylar and chalazal end wall and lateral wall of the embryo sac become more massive during endosperm development. Wall ingrowths also occur on the outer walls of the outer layer endosperm cells at both ends and lateral region of the embryo sac. When the embryo matures, many layers of chalazal endosperm ceils including 2–4 layers of transfer cells, a few of micropylar endosperm cells and 1–5 layers of peripheral endosperm cells are present. The nutrients of the embryo and endosperm at different stages of development are also discussed.     

Development of Embryo and Endosperm and Nutrient Accumulation in Vigna sesquipedalis (L.) Fruwirth

The structure of embryo sac, fertilization and development of embryo and endosperm in Vigina sesquipedalis (L.) Fruwirth were investigated. Pollization occures 7–10h before anthesis, and fertilization is completed 10 h after anthesis. After fertilization, wall ingrowths are formed at the micropylar and chalazal ends of the embryo sac. Embryo development conforms to the Onagrad type, and passes through 2 or more celled proembryo, long stick-shaped, globular, heart shaped, torpedo, young embryo, growing and enlarging embryo and mature embryo. Wall ingrowths are formed on the walls of basal cells and outer walls of the cells at basal region of suspenser. The suspensor remains as the seed reaches maturity. The starch grains accumulate in the cells of cotyledons by 9–16 days after anthesis, and proteins accumulate by 12–18 days after. The endosperm development follows the nuclear type. The endosperm ceils form at the micropylar end, and remain free nuclear phase at chalazal end. The outer cells are transfer cells. Those cells at the micropylar end form folded cells with wall ingrowths. At heartembryo stage, the endosperm begins to degenerate and disintegrates before the embryo matures.     

New data about the suspensor of succulent angiosperms: Ultrastructure and cytochemical study of the embryo-suspensor of Sempervivum arachnoideum L. and Jovibarba sobolifera (Sims) Opiz

The development of the suspensor in two species ?? Sempervivum arachnoideum and Jovibarba sobolifera ?? was investigated using cytochemical methods, light and electron microscopy. Cytological processes of differentiation in the embryo-suspensor were compared with the development of embryo-proper. The mature differentiated suspensor consists of a large basal cell and three to four chalazal cells. The basal cell produces haustorial branched invading ovular tissues. The walls of the haustorium and the micropylar part of the basal cell form the wall ingrowths typical for a transfer cells. The ingrowths also partially cover the lateral wall and the chalazal wall separating the basal cell from the other embryo cells. The dense cytoplasm filling the basal cell is rich in: numerous polysomes lying free or covering rough endoplasmic reticulum (RER), active dictyosomes, microtubules, bundles of microfilaments, microbodies, mitochondria, plastids and lipid droplets. Cytochemical tests (including proteins, insoluble polysaccharides and lipids are distributed in the suspensor during different stages of embryo development) showed the presence of high amounts of macromolecules in the suspensor cells, particularly during the globular and heart-shaped phases of embryo development. The protein bodies and lipid droplets are the main storage products in the cells of the embryo-proper. The results of Auramine 0 indicate that a cuticular material is present only on the surface walls of the embryo-proper, but is absent from the suspensor cell wall. The ultrastructural features and cytochemical tests indicate that in the two species ?? S. arachnoideum and J. sobolifera ?? the embryo-suspensor is mainly involved in the absorption and transport of metabolites from the ovular tissues to the developing embryo-proper.     

Transfer aleurone cells inSetaria lutescens (Gramineae)

Summary Typical aleurone cells occur around the periphery of the caryopsis. These cells are tabular with moderately thick walls and lack cell wall ingrowths. Transfer aleurone cells only occur adjacent to the placental vascular bundle, which supplies the developing embryo and endosperm. These specialized aleurone cells are approximately columnar, with thick walls bearing ingrowths on the outer radial and outer tangential walls. The wall ingrowths of transfer aleurone cells appear similar to those of transfer cells previously described and quite likely also function in short-distance transport of substances.Journal paper No. J-6737 of the Iowa Agricultural and Home Economics Experiment Station, Ames, Iowa. Project No. 1685.     

西瓜胚乳衰退过程中的超微结构变化及其对胚发育的作用

西瓜胚乳细胞衰退过程中 ,质膜、液泡膜突起、形成体积较大的囊泡 ,内质网断裂形成体积较小的囊泡 ;细胞质和细胞核降解形成电子致密的碎片沿细胞壁分布 ;细胞壁在衰退过程逐渐变薄 ,由于部分区域分解而使整个壁呈波浪型 ,细胞降解后的物质可直接穿越薄壁处或通过宽约 5 0 nm的胞间连丝向近胚端的胚乳细胞转移。胚乳与珠心组织分界壁 -胚囊壁上有发达的壁内突 ,有利于珠心组织内的物质向胚乳内转运 ;胚乳发育早期与胚共有的壁上内外两侧均有胼胝质沉积 ,壁上无外连丝型的胞间连丝存在 ,胚乳发育后期共有壁上的胼胝质消失 ,胚乳细胞降解物可穿越共有壁进入胚细胞内。实验结果表明西瓜胚乳在发育后期对胚的发育具有重要的作用。     

Ultrastructure of endopolyploid antipodals inAconitum vulparia Rchb.

Summary The ultrastructure of antipodals ofAconitum vulparia Rchb. was studied at two stages of development: at the earlier stage the endosperm has several nuclei, at the later one the endosperm is multinucleate. Over the investigated period the antipodal size enlarges distinctly. The wall ingrowths increase in size and number. Finally, they occur throughout the antipodal walls except for a small area in the extreme chalazal part, sunk deep into the nucellar podium. There are no plasmodesmata in the antipodal cell walls. The cytoplasm is dense and rich in ribosomes; it shows weak vacuolation. The rough endoplasmic reticulum is well developed. At the later stage dilated cisternae of endoplasmic reticulum are formed. Mitochondria, plastids and active dictyosomes are abundant. At the later stage some giant mitochondria are present; their matrix contains a large clear area with fine fibrils and an aggregation of fibrillar material. At this stage of development plastids have two types of inclusions: electron-transparent vacuoles and aggregations of electron-dense granules. The giant endopolyploid nuclei are considerably larger than those at the mature embryo sac stage; they are lobed on all sides.During the studied periodA. vulparia antipodals seem to be at their most active state.     

Structure of Embryo Sac Before and After Fertilization and Distribution of Transfer Cells in Ovules of Green Gram

The structure of embryo sac before and after fertilization, embryo and endosperm development and transfer cell distribution in Phaseolus radiatus were investigated using light and transmission electron microscopy. The synergids with distinct filiform apparatus have a chalazal vacuole, numerous mitochondria and ribosomes. A cell wall exists only around the micropylar half of the synergids. The egg cell has a chalazally located nucleus, a large micropylar vacuole and several small vacuoles. Mitochondria and plasrids with starch grains are abundant. No cell wall is present at its chalazal end. There are no plasma membranes between the egg and central cell in several places. The zygote has a complete cell wall, abundant mitochondria and plastids containing starch grains. Both degenerated and persistent synergids migh.t serve as a nutrient supplement to proembryo. The wall ingrowths occur in the central cell, basal cell, inner integumentary cells, suspensor cells and endosperm cells. These transfer cells may contribute to embryo nutrition at different developmental stages of embryo.     

Cytokinesis in the developing wheat grain; Division with and without a phragmoplast

Summary Cell wall formation during the transition from free-nuclear to cellular endosperm of wheat (Triticum aestivum L. cv. Heron) was investigated using correlated light and electron microscopy. Partitioning of the multinucleate syncytium that lines the inner periphery of the embryo sac is initiated 1–2 days after anthesis. Wall ingrowths, at first recognizable in sections as minute wall pegs, furrow inward from the edge of the embryo sac through the vacuolate cytoplasm which, to the inside, is clearly delimited by the central vacuole. Growth of the walls at this stage is independent of a phragmoplast and in this respect is reminiscent of the cleavage processes of lower plant cells. Intense fluorescence of the walls after staining with aniline blue suggests that callose may be a principal component. The growing walls branch and eventually meet on the side nearest the central vacuole. Cellularization of the peripheral layer of endosperm cytoplasm is thus complete about 2 days after anthesis. Between 2 and 3 days after anthesis, the peripheral layer of cells commences to divide both radially and tangentially and by 4 days the entire embryo sac is cellular. Cytokinesis during this phase entails the formation of a cell plate between sister nuclei. At the periphery of a forming cell plate, vesicles appear scattered amongst an array of phragmoplast microtubules. This mechanism of wall growth differs markedly from the initial infurrowing of the first-formed walls. The overall timing and the manner of cell wall deposition vary in a number of important respects from the model recently proposed by Mares et al. whose work was based largely on light microscopy (D.J. Mares; K. Norstog; A.B. Stone: Aust. J. Bot. 23, 311–326, 1975).Abbreviations CV central vacuole - D dictyosome - En endosperm - ER endoplasmic reticulum - II inner integument - m mitochondrion - MTs microtubules - N nucellus - NE nucellar epidermis - Nu nucleus - S starch - V vacuole - W embryo sac wall This work was supported by grants from the Australian Research Grants Committee and the Reserve Bank of Australia (to T.P. O'B.) while one of us (I.N.M.) received financial assistance from the Australian Government through the Commonwealth Scholarship and Fellowship Plan.     

Microtubular configurations during the cellularization of coenocytic endosperm in Ranunculus sceleratus L.

Summary Endosperm cellularization in Ranunculus sceleratus was studied in terms of the initiation of cell-wall formation in the coenocytic endosperm. The first endosperm cell walls were in an anticlinal position relative to the cell wall of the embryo sac and originated from the cell plates and not from wall ingrowths from the embryo-sac wall itself. Alveolar endosperm was formed 3 days after pollination. Microtubules were associated with the freely growing wall ends of the anticlinal walls and were observed in various orientations that generally ranged from angles of 45 ° to 90 ° to the plane of the wall. They were absent in the regions where vesicles had already fused. These microtubules may function in maintaining the growth and the direction of growth of the anticlinal wall until cellularization is completed. At the site where three neighbouring alveoli share their freely growing wall ends, remarkable configurations of microtubules were observed: in each alveolus, microtubules ran predominantly parallel to the bisector of the angle formed by the common walls. These microtubules may form a physically stable framework and maintain the direction of growth of the wall edges. It is concluded that the growing edge of the anticlinal endosperm wall and its associated microtubules are a special continuum of the original phragmoplast that gave rise to the anticlinal wall.     

The Initiation, Development and Removal of Embryo Sac Wall Ingrowths in the Developing Seeds of Solanum nigrum L. An Ultrastructural Study

In developing seeds of Solanum nigrum L., wall ingrowths developedat the extreme micropylar and chalazal ends of the embryo sac.In the micropylar region, the wall ingrowths were initiatedat the three-celled endosperm stage starting at the base ofthe zygote then progressing for a short distance chalazalwards.They developed quickly with the most elaborate around the baseof the suspensor. The chalazal wall ingrowths developed alongthe surfaces of the chalazal cup, the antipodal cup and thehypostase. Those along the hypostase were initiated at the four-celled,those in the chalazal and antipodal cups at the 20-celled endospermstages. The most elaborate developed along the base of the antipodalcup; the most simple were along the base of the chalazal cup.Small electron-lucent invaginations of the plasmalemma whichlater became filled with fibrillar material, were the earliestindication of wall ingrowth formation. Removal of the wall ingrowthscommenced at the mid-globular stage of embryo development andwas completed by the mid-heart-shaped stage. In the micropylarregion, wall ingrowth removal was rapid, starting with the lossof the fibrillar component followed by the thinning of the cellwall. However, along the hypostase and antipodal cup, a heterogeneouslayer of varying electron densities and a thinner, more electrondense layer was laid down over the ingrowths. This was followedby the removal of the fibrillar component. The initiation, removaland location of the embryo sac wall ingrowths is discussed inconnection with understanding the nutritional relationshipsbetween maternal tissue, endosperm and embryo.Copyright 1995,1999 Academic Press Wall ingrowths, Solanum nigrum, transfer cells, zone of separation and secretion, hypostase     

The specialized chalazal endosperm inArabidopsis thaliana andLepidium virginicum (Brassicaceae)

Summary Endosperm of the nuclear type initially develops into a large multinucleate syncytium that lines the central cell. This seemingly simple wall-less cytoplasm can, however, be highly differentiated. In developing seeds of members of the family Brassicaceae the curved postfertilization embryo sac comprises three chambers or developmental domains. The syncytium fills the micropylar chamber around the embryo, spreads as a thin peripheral layer surrounding a large central vacuole in the central chamber, and is organized into individual nodules and a large multinucleate cyst in the chalazal tip. Later in development, after the endosperm has cellularized in the micropylar and central chambers, the chalazal endosperm cyst remains syncytial and shows considerable internal differentiation. The chalazal endosperm cyst consists of a domelike apical region that is separated from the cellularized endosperm by a remnant of the central vacuole and a basal haustorial portion which penetrates the chalazal proliferative tissue atop the vascular supply. In the shallow chalazal depression ofArabidopsis thaliana, the cyst is mushroom-shaped with short tentacle-like processes penetrating the maternal tissues. The long narrow chalazal channel ofLepidium irginicum is filled by an elongate stalklike portion of the cyst. In both, the dome contains a labyrinth of endoplasmic reticulum, dictyosomes with associated vesicles, nuclei, and plastids. The basal portions, which lack the larger organelles, exhibit extensive wall ingrowths and contain parallel arrays of microtubules. The highly specialized ultrastructure of the chalazal endosperm cyst and its intimate association with degrading chalazal proliferative cells suggest an important role in loading of maternal resources into the developing seed.     

Early Endosperm Development in Ovules of Soybean, Glycine max (L) Merr. (Fabaceae)

Endosperm development was studied in normally setting flowersand pods of soybean from anthesis to a pod length of 10–20mm. The free-nuclear stage following double fertilization istypified by loss of starch and increasing vacuolation. The cytoplasmprovides evidence of extensive metabolic activity. Wall ingrowths,already present at the micropylar end of the embryo sac wallprior to fertilization, develop along the lateral wall of thecentral cell as well as at the chalazal endosperm haustorium.Endosperm cellularization begins when the embryo has developeda distinct globular embryo proper and suspensor. Cellularizationstarts at the micropylar end of the embryo sac as a series ofantidinal walls projecting into the endosperm cytoplasm fromthe wall of the central cell. The free, growing ends of thesewalls are associated with vesicles, microtubules, and endoplasrnicreticulum. Pendinal walls that complete the compartmentalizalionof portions of the endosperm cytoplasm are initiated as cellplates formed during continued mitosis of the endosperm nuclei.Endosperm cell walls are traversed by plasmodesmata. This studywill provide a basis for comparison with endosperin from soybeanflowers programmed to abscise. Glycine max, soybean, endosperm, ovules