Seed formation in two species of Adesmia (Fabaceae): co‐occurrence of micropylar and lateral endosperm haustoria in legumes and its taxonomic value

Seed ontogeny of Adesmia bicolor and Adesmia latifolia was analysed using light microscopy and standard histological techniques. Fertilization was porogamic. Linear proembryonal tetrads were observed in A. bicolor. The robust elongated suspensors possessed specialized basal cells. The nucellar epidermis became endothelial. The free‐nuclear endosperm produced a micropylar, filamentous and ephemeral haustorium and a lateral sac‐like haustorium at the funicular side. The cellular endosperm was initiated from the micropylar zone after the cordiform embryo stage. It mostly disintegrated in mature seeds. The sclerified bilayered testa was derived from the outer ovular integument. Different astrosclereid arrangements beyond the lens in both Adesmia species may be related to the different habitats of the two species. The occurrence of both micropylar and lateral nuclear endosperm haustoria has so far not been reported in Fabaceae and is the most distinctive embryological character of Adesmieae. The taxonomic value of the mostly uniform morphology of the suspensor in the Adesmia species studied could also be relevant. The nature of seed endothelia in many Fabaceae requires accurate redetermination prior to taxonomic use. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 158 , 602–612.     

Embryo sac, endosperm, and seed of Nemophila (Boraginaceae) relative to taxonomy, with a remark on embryogeny in Pholistoma

Studies on embryology and seed morphology are complementary to molecular phylogenetics and of special value at the genus level. This paper discusses the delimitation and evolutionary relationships of genera within the tribe Hydrophylleae of the Boraginaceae. The seven Nemophila species characterized by a conspicuous seed appendage are similar in embryology and seed structure. The ovule is tenuinucellate and unitegmic with a meristematic tapetum. The embryo sac penetrating the nucellar apex is of the Polygonum type, has short-lived antipodal cells, and an embryo sac haustorium. The endosperm is cellular, producing two terminal endosperm haustoria, of which the chalazal has a lateral branch. Embryogeny is of the Chenopodiad type (as in Pholistoma). The seed coat is formed from the small-celled inner epidermis of the integument. The large-celled outer epidermis of the integument disintegrates into scattered cells. Seed pits evolve from irregularly placed inner epidermal cells of the integument. The chalazal part of the ovule produces a cucullus, that functions as an ant-attracting elaiosome. Those species of Nemophila with a conspicuous cucullus form a natural genus. Nemophila is most closely related to Pholistoma. The integumentary seed pits of Nemophila might have evolved from ovular seed pits similar to those in Pholistoma.     

DEVELOPMENT OF THE SEED AND FRUIT IN RHINANTHUS MAJOR AND R. SEROTINUS

There are seven sessile, campylotropous, discoid ovules in each loculus of the anteroposteriorly flattened bilocular ovary. They are arranged alternately in two rows in each chamber on the axile placenta which is nodular where the ovules are borne. Nucellus degenerates early except at the chalazal end of the curved embryo sac, and the inntermost layer of the integument functions as endothelium. The aggressive, multinucleate micropylar haustorium grows as a tubular body through the micropylar canal and ramifies in the placenta while the two-nucleate chalazal haustorium creates a large space by digesting a good deal of the chalazal tissue. Endosperm is differentiated into three regions: the middle storage, the haustorial micropylar, and the chalazal. Thickness of the integument is considerably added to by the endothelium and by its surrounding meristematic zone of the integument. There are two prominent wings on the dorsal and smaller ones on the lateral faces of the cochlidiospermous seed, its ventral face being occupied by a prominent basal body. A heavily cutinized envelope, formed by the endothelium, surrounds the ovoid storage endosperm. Testa of the seed is mainly composed of the thickened epidermis and the endothelium. The micropylar and the chalazal parts of the endosperm become tanniferous and serve to plug the two ends of the seed. Embryo is straight, and it bears two cotyledons and two plumular leaves.     

Ephemeral and non-ephemeral structures during seed development in two Cytisus species (Papilionoideae,Leguminosae)

Abstract

Seed formation involves not only the embryo and endosperm development, but also the formation of a series of either ephemeral or non-ephemeral structures. In this article, we study several of those structures in Cytisus multiflorus and Cytisus striatus. The endosperm development is first nuclear and later cellular, except for the chalazal area, whose development is always nuclear. It generates, in the early developmental stages, a sac-like haustorium. As the seed develops, two structures seem to be closely related to nutrient mobilization to the embryo sac: on the one hand, a group of cells and a channel, located in the chalazal area and closely related between them and to the endosperm haustorium, which could be interpreted as a hypostase and on the other hand, an endothelium, derived from the inner integument, which later degenerates leaving no trace in the mature seed. All of these structures would be associated with the directionality of assimilates from ovule tissues to embryo sac. In mature seed and surrounding the embryo appears a unicellular layer of cells rich in proteins (aleurone layer), which is the origin of the outermost layer of the cellular endosperm. The seed coat is made up only of the outer integument.     

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。     

Development of megagametophyte, embryo, and seed in Senna corymbosa (Lam.) H.S. Irwin & Barneby (Leguminosae – Caesalpinioideae)

The main aspects of seed ontogeny in Senna corymbosa were studied by standard anatomical microtechniques for light microscope observations. The results revealed an ana-campylotropous, bitegmic, and crassinucelate mature ovule. A single archesporocyte developed by an archesporial cell enlargement from the subhypodermal multicellular archesporium. Meiosis originated linear or T-shaped megasporic tetrads. The functional megaspore was the chalazal one. Megagametophytic development conformed to the Polygonum type. Fertilization was porogamic. Endosperm development was free nuclear and conformed to a chalazal haustorium. Cellular endosperm was initiated from the micropylar end during the globular embryo stage. Embryogeny derived from a linear proembryonal tetrad. The mature embryo showed an oblique axis. The testa derived from the outer ovular integument. Nucellar and endosperm remnants, and the micropylar region of the inner ovular integument, persisted at embryo maturity. The absence of a pleurogram would be adaptative to wetland habitats. The taxonomic use of the mature embryo axis in the Cassieae and the phylogenetic employment of megasporic arrangements in Leguminosae needs some reinterpretation.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 153 , 169–179.     

Le Basi Anatomiche Dei Rapporti Nutritivi Tra Pianta Madre,Seme Ed Embrione

Abstract

The anatomical basis of the nutritive relationships between mother plant, seed, and embryo. — The morphology and anatomy of the fruits and seeds of the Angiosperms show a great variety of structures and adaptments, even within the same family, and one must be cautious in drawing generalized conclusions.

If we first examine the ovary we see that the single carpel receives three vascular traces from which the three main bundles originate, a dorsal and two ventral ones, all more or less reduced. Except in the case of laminar placentation the ovule traces are connected to the ventral vascular system, but often the entire vascular system of the ovary is anastomosed and therefore reticular. However especially when the placentae are at the centre or at the basis of the ovary, it is possible to detect a tendency towards a separation between the vascular system of the ovarian wall and that of the placentae.

The ovular bundle runs through the funicle reaching the chalaza, where it can either end or continue towards the micropyle with a single bundle or with a few branches or even forming a complete reticular envelope surrounding the ovule. The ovular vascular bundles are normally found in the outer integument.

The ovule is made of an inner part (nucellus), and an outer one (integuments). The integuments play a very important role in the processes of seed maturation, dormancy, and germination. They are isolated from the interior of the seed by a cuticle which is a common production of the inner integumentary epidermis, and of the nucellus. The cuticle is not present in the chalaza and can be dissolved in the micropylar region: through these two apertures nutrients can penetrate into the seed or haustoria can grow out of it. During the course of maturation these openings become closed by various means, often through the formation of a new cuticle or of a suberised chalazal plate.

The nutrients which pass through the chalaza penetrate into the nucellus where in some cases one can find some structures which facilitate the communications between the chalaza and embryo sac. The endosperm feeds at the expense of the nucellus but often it can establish a direct contact with the chalaza or the integuments or even the placentae. This occurs often thanks to haustoria.

The embryo is normally surrounded at first by a more or less liquid endosperm: in a second stage the endosperm becomes cellular and the embryo grows at its expense through the digestive activity of the cotyledonar epidermis.

From an anatomo-physiological point of view the following points seem of particular interest:

(I) The endosperm and the embryo show a remarkable autonomy in respect of the mother plant: from an anatomical point of view this is shown by the isolation of the endosperm and embryo by means of a cuticular covering or substitutive structures and by the interposition of nutritive tissues between the vascular system of the mother plant and the endosperm.

(II) Given the importance of the inner cuticle its presence and its constitution should be ascertained in the various species having also in mind the properties of selective permeability shown by the testa.

(III) Two nutritive mechanisms exist: translocation of nutrients via the vascular system and the nutritive tissues, and digestion of surrounding cells. In the digestive phenomena it is important to explain the mechanisms by which only the right cells are digested and not the others.

(IV) The embryo very frequently is immersed at first in a more or less liquid endosperm and is later surrounded by a compact tissue; the nutritive mechanisms are probably different in the two cases.

(V) Two endospermic zones are often distinguishable: one having an haustorial or at least a digestive or elaborative function, and being typically non cellular; another zone, typically cellular, forms a tissue which is sooner or later absorbed by the embryo. The cellularization of this zone seems to coincide with the establishment of polarity and with the beginning of maximum growth of the embryo.

(VI) The relationships between the inner seed and the integuments is complex and there is a correlation between the histoanatomical and biochemical changes of these two parts during seed development. The modifications undergone by the integuments are important steps also towards the preparation of the seed to the processes of dispersal, dormancy, and germination.     

黄精种子萌发过程发育解剖学研究

采用石蜡切片技术对成熟黄精种子形态及萌发过程中的形态学变化及解剖结构特征进行了研究,以阐明黄精种子繁殖的生物学机制。结果显示:(1)成熟的黄精种子由外而内依次为种皮、胚乳和胚等3部分组成。其中种皮由一层木质化的细胞组成;胚乳占据种子的大部分结构,胚乳细胞含有大量淀粉,细胞壁增厚;胚处于棒型胚阶段。(2)黄精种子在萌发过程中棒型胚靠近种脐端分化为吸器、子叶联结和子叶鞘,靠近种孔的部位分化出胚根、胚轴和胚芽。(3)黄精种子萌发首先由子叶联结伸长将胚芽和胚根原基推出种孔,紧接着下胚轴膨大形成初生小根茎,吸器留在种子中分解吸收胚乳中的营养物质。(4)通过子叶联结连通吸器和初生小根茎,将胚乳中的营养物质由吸器-子叶联结这个通路转移到初生小根茎中,为初生根茎上胚芽和胚根的进一步分化提供物质保障。(5)黄精种子自然条件下萌发率较低,而且当年不出土。研究表明,黄精种子的繁殖生物学特性是其生态适应的一种重要机制。     

THE DEVELOPMENT OF THE SEED IN ANDROGRAPHIS SERPYLLIFOLIA

Mohan Ram , H. Y. (U. Delhi, India.) The development of the seed in Andrographis serpyllifolia. Amer. Jour. Bot. 47(3) : 215—219. Illus. 1960.–Andrographis serpyllifolia, a member of the Acanthaceae, has an embryo sac with a bifurcated chalazal part. At the time of fertilization both synergids and antipodal cells disintegrate. Early in its development the endosperm is composed of 3 distinct parts: (1) a binucleate densely cytoplasmic chalazal haustorium; (2) a large binucleate micropylar haustorium; and (3) a central chamber which develops into the endosperm proper. The divisions in the central endosperm chamber are ab initio cellular. A few of the endosperm cells elongate enormously, ramify into the integument and destroy the surrounding cells. These cells have been termed secondary haustoria. Due to the unequal destruction of the integument, the endosperm assumes a ruminate condition. The mature seed is nearly naked because the seed coat is almost completely digested. The embryo has a long suspensor. The micropylar cells of the suspensor are hypertrophied and multinucleate. Contrary to Mauritzon's (1934) belief, the course of endosperm development is markedly different from that observed in Thunbergia. So far, albuminous seeds have been reported only in the subfamily Nelsonioideae. The present investigation provides a case of its occurrence in the Acanthoideae also.     

罗汉果胚、胚乳及胚乳吸器的发育

本文采用石蜡切片与酶解分离法对罗汉果Ｓｉｒａｉｔｉａｇｒｏｓｖｅｎｏｒｉ胚、胚乳及胚乳吸器的发育过程进行观察．ａ）罗汉果胚的发育是按Ｇｅｕｍｕｒｂａｎｕｍ的分裂程序进行的．属紫菀型．但在合子分裂成球胚过程中,胚芽原细胞分化明显．故属紫菀型的变异型。ｂ）胚乳发育属核型．在球形胚阶段,在合点端和珠孔端有发育的胚乳吸器形成并进行旺盛生长,最大长度达１４２０μｍ,心形胚期．吸器活动开始减退,合点端核型胚乳吸器转变成细胞型．由胚乳本体基部膨大细胞．充当补助吸器．ｃ）酶解分离法研究胚乳吸器的发生发育有较好的应用前景。     

Embryological features of Alhagi persarum (Fabaceae): Adapt to environmental constraints

Reproductive organs, in flowering plants, are sensitive to stressful environments. Alhagi persarum Boiss. & Buhse copes with the stresses and produce reproductive organs under difficult climatic conditions. Embryological characters of this plant were investigated for the first time using different microscopy and staining techniques. The results of this study showed unique reproductive characters and strategies in A. persarum that we named reproductive adaptation. These characters have roles in protection and nutrition of reproductive organs, some of which were visible in ovule: accumulation of phenolic compounds, presence of ovular endothelium with its cuticle coat, hypostase, postament, endosperm haustorium, presence of operculum, curvature of the embryonic axis. The other characters in the seed are macrosclereid cells with cuticle coat, double palisade layer and lignified tracheids in hilar groove. Thickness increasing of endothecium and exine are the adaptive characters in anther. Unlike many of the stress-sensitive plants, all developmental stages of the embryo sac, anther, pollen and pollen tube are without any defects in these stress-tolerant plants. Seed germination rate is low in this species that is due to the hardness of seed coat which causes seed deep exogenous dormancy. This dormancy is also a developmental program for stress tolerance to keep seed viability for a long time in difficult conditions.     

西瓜胚和胚乳的发育

应用显微技术对西瓜胚和胚乳的发育过程进行了观察并分析了西瓜胚珠败育的原因。西瓜胚发育属紫菀型。合子第一次分裂为不均等分裂 ,形成的基细胞体积明显较顶细胞大 ,两细胞均含有多个液泡。原胚发育过程中没有明显的胚柄。最外层的原胚细胞 ,与胚乳细胞相邻的壁上被胼胝质物质包围 ,且无外连丝存在 ;与胚囊壁相接的壁上无壁内突结构。胚的子叶体积增长的同时 ,子叶细胞内积累蛋白质和脂类物质 ,多糖物质的含量下降。胚乳发育属核型 ,在球形胚期开始自珠孔端向合点端细胞化 ,胚子叶分化出后开始自珠孔端向合点端退化。胚乳合点端在球形胚早期形成发达的胚乳吸器 ,开始呈游离核状态 ,后细胞化 ,在心型胚期之后退化。     

PATTERNS OF EMBRYO AND ENDOSPERM FORMATION IN MONOTROPA HYPOPITYS (MONOTROPACEAE) FROM NORTH AMERICA AND WESTERN SWEDEN

Within the Monotropaceae. Monotropa hypopitys L. has the widest geographical distribution with sporophytes characterized as achlorophyllous, mycotrophic, and morphologically reduced. General and histochemical observations at the light microscope level concerning the postpollination changes in the numerous anatropous, unitegmic ovules reveal a precise embryogeny and endosperm development. Following double fertilization, the primary endosperm cell produces a lipid-rich cellular endosperm situated between a micropylar and a chalazal haustorium. A cytoplasmically unequal division of the elongated zygote initiates proembryo formation. The degeneration of the basal cell of the proembryo results in an isolated terminal cell that undergoes a cytoplasmically equal, transverse division establishing a two-celled embryo embedded in endosperm. Prior to final seed maturation, proteins replace the lipids as the dominant cytoplasmic reserve material. In contrast with earlier studies that depicted the mature embryo as variable in structure, here the embryo is shown to be consistently uniform within and between those populations sampled from North America and Europe.     

Occurrence of Endosperm Haustorium in Cannabis sativa L.: With thirteen Figures in the Text

The development and structure of the chalazal endosperm haustoriumin Cannabis sativa are described. The endosperm is nuclear anda haustorium is formed at the chalazal end. The latter remainsfree nuclear throughout. Enucleate vesicles appear in the upperpart of the endosperm but finally they merge with the cytoplasmof the haustorium. As the embryo reaches maturity it occupiesthe whole seed cavity, the haustorium collapses and the endospermpersists only as a thin layer.     

西瓜胚乳吸器的发育及ATP酶的超微细胞化学定位

报道了西瓜（Citrullus lanatus)胚乳吸器发育过程,并对胚乳吸器细胞中的ATP酶进行了超微细胞化学定位,球形胚早期,胚囊合点端的壁伸长发育成一管状胚乳吸器,进而吸器靠近乳本体端膨大为囊状,球形胚晚期吸器自珠孔端向合点端逐渐细胞化,胚分化出子叶时,胚乳吸器自合点端向珠孔端退化,在刚形成的胚乳吸器细胞中,ATP酶活性反应主要分布在细胞的核膜,内质网上,胞间连丝和吸器细胞壁内的小球状物上也有较强的ATP酶活性反应;在开始退化的吸器细胞中,核膜上的ATP酶性的反应减弱较早,内质网稍晚,进一步退化的胚乳吸器细胞中,ATP酶主要集中分布在细胞壁,细胞间隙内,核上几乎没有ATP酶性反应,内质网上仅有微弱的ATP酶反应。     

Organisation of the endosperm and endosperm–placenta syncytia in bladderworts (Utricularia, Lentibulariaceae) with emphasis on the microtubule arrangement

Multinucleate cells play an important role in higher plants, especially during reproduction; however, the configurations of their cytoskeletons, which are formed as a result of mitosis without cytokinesis, have mainly been studied in coenocytes. Previous authors have proposed that in spite of their developmental origin (cell fusion or mitosis without cytokinesis), in multinucleate plant cells, radiating microtubules determine the regular spacing of individual nuclei. However, with the exception of specific syncytia induced by parasitic nematodes, there is no information about the microtubular cytoskeleton in plant heterokaryotic syncytia, i.e. when the nuclei of fused cells come from different cell pools. In this paper, we describe the arrangement of microtubules in the endosperm and special endosperm–placenta syncytia in two Utricularia species. These syncytia arise from different progenitor cells, i.e. cells of the maternal sporophytic nutritive tissue and the micropylar endosperm haustorium (both maternal and paternal genetic material). The development of the endosperm in the two species studied was very similar. We describe microtubule configurations in the three functional endosperm domains: the micropylar syncytium, the endosperm proper and the chalazal haustorium. In contrast to plant syncytia that are induced by parasitic nematodes, the syncytia of Utricularia had an extensive microtubular network. Within each syncytium, two giant nuclei, coming from endosperm cells, were surrounded by a three-dimensional cage of microtubules, which formed a huge cytoplasmic domain. At the periphery of the syncytium, where new protoplasts of the nutritive cells join the syncytium, the microtubules formed a network which surrounded small nuclei from nutritive tissue cells and were also distributed through the cytoplasm. Thus, in the Utricularia syncytium, there were different sized cytoplasmic domains, whose architecture depended on the source and size of the nuclei. The endosperm proper was isolated from maternal (ovule) tissues by a cuticle layer, so the syncytium and chalazal haustorium were the only way for nutrients to be transported from the maternal tissue towards the developing embryo.     

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。     

Effects of APETALA2 on embryo, endosperm, and seed coat development determine seed size in Arabidopsis

Arabidopsis APETALA2 (AP2) controls seed mass maternally, with ap2 mutants producing larger seeds than wild type. Here, we show that AP2 influences development of the three major seed compartments: embryo, endosperm, and seed coat. AP2 appears to have a significant effect on endosperm development. ap2 mutant seeds undergo an extended period of rapid endosperm growth early in development relative to wild type. This early expanded growth period in ap2 seeds is associated with delayed endosperm cellularization and overgrowth of the endosperm central vacuole. The subsequent period of moderate endosperm growth is also extended in ap2 seeds largely due to persistent cell divisions at the endosperm periphery. The effect of AP2 on endosperm development is mediated by different mechanisms than parent-of-origin effects on seed size observed in interploidy crosses. Seed coat development is affected; integument cells of ap2 mutants are more elongated than wild type. We conclude that endosperm overgrowth and/or integument cell elongation create a larger postfertilization embryo sac into which the ap2 embryo can grow. Morphological development of the embryo is initially delayed in ap2 compared with wild-type seeds, but ap2 embryos become larger than wild type after the bent-cotyledon stage of development. ap2 embryos are able to fill the enlarged postfertilization embryo sac, because they undergo extended periods of cell proliferation and seed filling. We discuss potential mechanisms by which maternally acting AP2 influences development of the zygotic embryo and endosperm to repress seed size.     

Seed fertilization, development, and germination in Hydatellaceae (Nymphaeales): Implications for endosperm evolution in early angiosperms

New data on endosperm development in the early-divergent angiosperm Trithuria (Hydatellaceae) indicate that double fertilization results in formation of cellularized micropylar and unicellular chalazal domains with contrasting ontogenetic trajectories, as in waterlilies. The micropylar domain ultimately forms the cellular endosperm in the dispersed seed. The chalazal domain forms a single-celled haustorium with a large nucleus; this haustorium ultimately degenerates to form a space in the dispersed seed, similar to the chalazal endosperm haustorium of waterlilies. The endosperm condition in Trithuria and waterlilies resembles the helobial condition that characterizes some monocots, but contrasts with Amborella and Illicium, in which most of the mature endosperm is formed from the chalazal domain. The precise location of the primary endosperm nucleus governs the relative sizes of the chalazal and micropylar domains, but not their subsequent developmental trajectories. The unusual tissue layer surrounding the bilobed cotyledonary sheath in seedlings of some species of Trithuria is a belt of persistent endosperm, comparable with that of some other early-divergent angiosperms with a well-developed perisperm, such as Saururaceae and Piperaceae. The endosperm of Trithuria is limited in size and storage capacity but relatively persistent.     

DEVELOPMENT OF NORMAL AND SEEDLESS ACHENES IN CICHORIUM INTYBUS (COMPOSITAE)

Cross- and partially cross-pollinated capitula of Cichorium intybus (Compositae, Lactuceae) were examined for a study of normal and seedless fruit development respectively. Embryos develop according to the Asterad pattern, and the free-nuclear endosperm becomes cellular 15–17 hrs after pollination. A zone of disorganized cellular material surrounds the embryo sac at anthesis, and, in normal achenes, this zone expands as the seed develops. Initially the developing seed elongates and comes into contact with the top of the ovary by 48 hrs. In contrast to this pattern, the ovule in developing seedless achenes degenerates within 72 hrs. Irregularities, such as an abnormally proliferating endothelium, embryo formation without endosperm, and endosperm formation without an embryo often accompany this degeneration. Differentiation of the pericarp in seeded achenes begins between 48 and 72 hrs, starting at the apex and proceeding basipetally; in seedless fruits the process is similar though initiated somewhat later. The normal pericarp at maturity exhibits a pigmented exocarp, a broad mesocarp of thick-walled lignified cells, and a tenuous endocarp. In seedless achenes the fruit coat is similar except that the exocarp is colorless and the cells of the mesocarp are relatively small.