Embryology in Trithuria submersa (Hydatellaceae) and relationships between embryo, endosperm, and perisperm in early-diverging flowering plants

? Premise of the study: Despite their highly reduced morphology, Hydatellaceae bear the unmistakable embryological signature of Nymphaeales, including a starch-rich maternal perisperm and a minute biparental endosperm and embryo. The co-occurrence of perisperm and endosperm in Nymphaeales and other lineages of flowering plants, and their respective functions during the course of seed development and embryo germination, remain enigmatic. ? Methods: Development of the embryo, endosperm, and perisperm was examined histologically from fertilization through germination in flowers and fruits of Trithuria submersa. ? Key results: The embryo of T. submersa initiates two cotyledons prior to seed maturity/dormancy, and their tips remain in contact with the endosperm throughout germination. The endosperm persists as a single layer of cells and serves as the interface between the embryo and the perisperm. The perisperm contains carbohydrates and proteins, and functions as the main storage tissue. The endosperm accumulates proteins and aleurone grains and functions as a transfer cell layer. ? Conclusions: In Nymphaeales, the multiple roles of a more typical endosperm have been separated into two different tissues and genetic entities: a maternal perisperm (nutrient acquisition, storage, mobilization) and a minute biparental endosperm (nutrient transfer to the embryo). The presence of perisperms among several other ancient lineages of angiosperms suggests a modest degree of developmental and functional lability for the nutrient storage tissue (perisperm or endosperm) within seeds during the early evolution of flowering plants. Finally, we examine the evolutionary developmental hypothesis that, contrary to longstanding assumptions, an embryo-nourishing perisperm along with a minute endosperm may represent the plesiomorphic condition for flowering plants.     

象牙参种子的解剖学和组织化学研究

象牙参种子解剖学和组织化学的研究结果表明, 种子包括假种皮、种皮、外胚乳、内胚乳和胚。假种皮没有完全包被种子, 由约4~5 层薄壁细胞构成。种皮可以分为外种皮、中种皮和内种皮。外种皮由1 层表皮细胞构成, 细胞壁明显增厚;中种皮包括下皮层、半透明细胞层和3~4层细胞的色素层, 下皮层和色素层细胞均充满红棕色色素;内种皮由1 层体积小、壁局部增厚的砖形薄壁细胞构成。种子在珠孔端分化出珠孔领、孔盖和种阜状结构, 珠孔领为同形型, 孔盖不具石细胞硬层。合点区内种皮出现缺口, 缺口间充满合点区色素细胞, 其整体轮廓成新月形。外胚乳可分为厚区与薄区两部分, 外胚乳细胞壁平直, 细胞内充满淀粉。内胚乳细胞主要含蛋白质, 也有少量脂类物质, 细胞界限不清楚。胚棒状, 两端略膨大, 含大量脂类物质, 也含蛋白质和多糖。     

拟豆寇种子的解剖学和组织化学研究

拟豆寇（Paramomum petaloideum)种子包括假种皮,种皮,外胚乳,内胚乳和胚,假种皮膜质,由5－7层薄壁细胞构成,种皮分外种皮,中种皮和内种皮,外种皮由一层细胞构成,其壁增厚并木质化,中种皮包括下皮导,透明细胞层和色素层,下皮由一层细胞构成,细胞近长方形,半透明细胞层由一层细胞组成,细胞近长方形或长条形,形态上与色素层细胞相似,但可通过染色方法把二者区分,色素层由一至二层细胞构成,最内层细胞形态有时难以分辨,内种皮由一层内切几壁非常增厚的石细胞构成,珠孔区分化出珠孔领,孔盖和珠孔区薄壁细胞,珠孔领导形型,孔盖具有石细胞硬具,合点区内种皮内凹陷并出现缺口,缺口位于种子近顶部偏向背侧,缺口间的合点区色素细胞群整体轮廓呈喇形,壁呈波浪形的外胚细胞富含淀粉粒,内胚乳最外一层细胞体积小,富含脂类物质,内胚乳合点端多层细胞,珠孔端为一层细胞,含丰富蛋白质,脂类物质主要存在于胚中,本文还从种子解剖学角度讨论了拟豆寇的系统位置。     

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.     

Ovule morphology, embryogenesis and seed development in three Hylocereus species (Cactaceae)

Pre-embryonic and embryonic stages and seed developments were studied in the diploids Hylocereus monacanthus and Hylocereus undatus and the tetraploid Hylocereus megalanthus. Ovule morphology was similar among species except for micropyle entrance. H. monacanthus had the thickest and most robust suspensor. Embryo developmental time, measured from fertilization to maturity, was significantly more prolonged in H. megalanthus. Typical to Cactaceae, the seed coat was formed by one layer of sclerenchymatous cells, but was more lignified in H. megalanthus. Morphological features common to all species included (1) cellular type endosperm with independent patterns of development in the chalazal and micropylar zones, forming a haustorium layer from the chalazal zone to the embryo; (2) an endothelial layer surrounding the embryo sac almost complete; (3) a nucellar summit growing into the micropyle; and (4) a placental obturator and a funicle connecting the ovarian tissue to the ovule. Seed development was typically endospermic (exendospermic orthodox seeds). Anomalies included two egg cells in the same embryo sac, two embryos developing in the same ovule, and embryos developing from the chalazal pole region. Total seed number and seed viability were significantly lower in H. megalanthus than in the other two taxa. Embryos at different developmental stages were observed in aborted H. megalanthus seeds.     

ADVENTIVE EMBRYOGENESIS IN CITRUS (RUTACEAE) II. POSTFERTILIZATION DEVELOPMENT

Cytological and histological studies on postfertilization development of ovules were carried out in six facultatively apomictic Citrus cultivars. At the time of anthesis, adventive embryo initial cells (AEICs) were detected mainly in the cell layers of the nucellus around the chalazal half of the embryo sac. During the approximately 40 days rest period of the AEICs after fertilization, rapid cell division and enlargement in the endosperm and the chalazal half of the nucellus resulted in the split of AEICs into several separated areas forming the micropylar, lateral and chalazal islands surrounding the enlarging embryo sac. Both in diploid seeds with triploid endosperm and triploid seeds with pentaploid endosperm, the AEICs located in the micropylar half successfully developed into adventive embryos. In diploid seeds, almost all AEICs located in the chalazal half did not develop beyond the initial-celled stage, while in the triploid seeds, those located in the chalazal half occasionally developed into cotyledonary embryos. In seeds with aborted endosperm, the AEICs located in the chalazal half often developed into cotyledonary embryos. The chalazal expiants from normal seeds produced a large number of embryos in vitro. Four results can be summarized from these studies on adventive embryogenesis as follows: 1) All AEICs are initiated prior to anthesis. 2) Whether or not the AEICs successfully developed into adventive embryos is dependent upon their position in the seed. 3) The farther the AEICs are located from the micropylar end, the more adventive embryogenesis is suppressed by endosperm. 4) The degree of adventive embryogenesis in the chalazal half is affected by time and extent of malfunction of the endosperm. Under natural conditions, these regulatory systems of adventive embryogenesis contribute to high production of zygotic seedlings in apomictic Citrus species and cultivars.     

Seed ultrastructure and water absorption pathway of the root-parasitic plant Phelipanche aegyptiaca (Orobanchaceae)

Background and Aims

Obligate root parasitic plants of the Orobanchaceae do not germinate unless they chemically detect a host plant nearby. Members of this family, like Orobanche, Phelipanche and Striga, are noxious weeds that cause heavy damage to agriculture. In spite of their economic impact, only a few light microscopical studies of their minute seeds have been published, and there is no knowledge of their ultrastructure and of the role each tissue plays during the steps preceding germination. This paper describes the ultrastructure of Phelipanche seeds and contributes to our understanding of seed tissue function.

Methods

Seeds of P. aegyptiaca were examined under light, scanning electron, transmission electron and fluorescence microscopy following various fixations and staining protocols. The results were interpreted with physiological data regarding mode of water absorption and germination stimulation.

Key Results and Conclusions

The endothelium, which is the inner layer of the testa, rapidly absorbs water. Its interconnected cells are filled with mucilage and contain labyrinthine walls, facilitating water accumulation for germination that starts after receiving germination stimuli. Swelling of the endothelium leads to opening of the micropyle. The perisperm cells underneath this opening mediate between the rhizosphere and the embryo and are likely to be the location for the receptors of germination stimuli. The other perisperm cells are loaded with lipids and protein bodies, as are the endosperm and parts of the embryo. In the endosperm, the oil bodies fuse with each other while they are intact in the embryo and perisperm. Plasmodesmata connect the perisperm cells to each other, and the cells near the micropyle tightly surround the emerging seedling. These perisperm cells, and also the proximal embryo cells, have dense cytoplasmic contents, and they seem to represent the two seed components that are actively involved in transfer of reserve nutrients to the developing seedling during germination.     

Seed-specific expression of seven Arabidopsis promoters

Seeds contain storage compounds, from various carbohydrates to proteins and lipids, which are synthesized during seed development. For the purposes of many plant researches or commercial applications, developing promoter systems expressing specifically in seeds or in particular constituents or tissues/compartments of seeds are indispensable. To screen genes dominantly or specifically expressed in seed tissues, we analyzed Arabidopsis ATH1 microarray data open to the public. Thirty-two candidate genes were selected and their expressions in seed tissues were confirmed by RT-PCR. Finally, seven genes were selected for promoter analysis. The promoters of seven genes were cloned into pBI101 vector and transformed into Arabidopsis to assay histochemical β-glucuronidase (GUS) activity. We found that Pro-at3g03230 promoter drove GUS expression in a chalazal endosperm, Pro-at4g27530:GUS expressed in both chalazal endosperm and embryo, Pro-at4g31830 accelerated GUS expression both in radicle and procambium, Pro-at5g10120 and Pro-at5g16460 drove GUS expression uniquely in embryo, Pro-at5g53100:GUS expressed only in endosperm, and Pro-at5g54000 promoted GUS expression in both embryo and inner integument. These promoters can be used for expressing any genes in specific seed tissues for practical application.     

距药姜种子解剖学和组织化学研究

距药姜种子解剖学和组织化学研究表明,种子包括种皮、外胚乳、内胚乳和胚。外皮由1层表皮细胞构成,细胞壁纤维素质且明显增厚,中种皮可分为1层细胞的下皮层、半透明细胞层和2-4层细胞的色素细胞层,下皮层和色素细胞层的细胞内充满棕红色色素;内种皮由1层砖形薄壁细胞构成。珠孔区有珠孔领和孔盖的分化,但珠孔领分化不完善。合点区内种皮出现缺口,缺口间充满合点区色素细胞,其整体轮廓成新月形。外胚乳细胞壁平直,细胞内充满淀粉。内胚乳可分为多细胞区简细胞区两部分,内胚乳细胞界限不清,内含物主要是蛋白质,胚少有分化,含脂类、蛋白质、多糖,另外,还对姜花族的种子解剖学特征进行了初步的系统学分析。     

Comparative ovule and megagametophyte development in Hydatellaceae and water lilies reveal a mosaic of features among the earliest angiosperms

Background and Aims: The embryo sac, nucellus and integuments of the early-divergentangiosperms Hydatellaceae and other Nymphaeales are comparedwith those of other seed plants, in order to evaluate the evolutionaryorigin of these characters in the angiosperms. Methods: Using light microscopy, ovule and embryo sac development aredescribed in five (of 12) species of Trithuria, the sole genusof Hydatellaceae, and compared with those of Cabombaceae andNymphaeaceae. Key Results: The ovule of Trithuria is bitegmic and tenuinucellate, ratherthan bitegmic and crassinucellate as in most other Nymphaeales.The seed is operculate and possesses a perisperm that developsprecociously, which are both key features of Nymphaeales. However,in the Indian species T. konkanensis, perisperm is relativelypoorly developed by the time of fertilization. Perisperm cellsin Trithuria become multinucleate during development, a featureobserved also in other Nymphaeales. The outer integument issemi-annular (‘hood-shaped’), as in Cabombaceaeand some Nymphaeaceae, in contrast to the annular (‘cap-shaped’)outer integument of some other Nymphaeaceae (e.g. Barclaya)and Amborella. The megagametophyte in Trithuria is monosporicand four-nucleate; at the two-nucleate stage both nuclei occurin the micropylar domain. Double megagametophytes were frequentlyobserved, probably developed from different megaspores of thesame tetrad. Indirect, but strong evidence is presented forapomictic embryo development in T. filamentosa. Conclusions: Most features of the ovule and embryo sac of Trithuria are consistentwith a close relationship with other Nymphaeales, especiallyCabombaceae. The frequent occurrence of double megagametophytesin the same ovule indicates a high degree of developmental flexibility,and could provide a clue to the evolutionary origin of the Polygonum-typeof angiosperm embryo sac.     

ADVENTIVE EMBRYOGENESIS IN CITRUS AND ITS RELATION TO POLLINATION AND FERTILIZATION

Cytological and histological studies of seeds from three facultative apomictic Citrus cultivars show that adventive embryos develop, as a rule, from the first few cell layers of the nucellus adjacent to the embryo sac in the micropylar half and occasionally from the chalazal end. The adventive embryos initiated in nucellar tissue away from the embryo sac and most of those initiated from the chalazal end of the nucellus do not develop beyond the one-celled stage. When two or more embryos are developing in the same seed, the successful development of a given embryo depends on its location in relation to access to nutrients from the endosperm. The presence of a zygote and triploid endosperm in seeds with adventive embryos, the abortion of seed when endosperm degenerates, and the lack of seed set without pollination indicate that pollination and fertilization are essential for in vivo adventive embryogenesis.     

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.     

姜目芭蕉群植物种子解剖学研究及其系统学意义

研究了姜目芭蕉群代表植物象腿蕉属象腿蕉(Ensete glaucum)、旅人蕉属旅人蕉(Ravenala madagascariensis)与蝎尾蕉属Heliconia faranmansis?D6肿咏馄侍卣鳌＝峁砻鳎笸冉段藜僦制ぃ制し只霰砥ぁ⒑癖谧橹赴褪赴悖赴瞿谇邢虮谟刖断虮谠龊瘢缓系闱哂泻系闶矣牒系愣眩谥制ち恢榭浊兄榭琢旌涂赘堑姆只榭琢煳涡停赘侵挥赡谥制は赴钩桑褐榭浊制ぱ由煨纬芍制昵唬和馀呷?层细胞：内胚乳细胞径向延长,细胞内充满淀粉粒。旅人蕉具假种皮,种皮分化出外种皮、中种皮和内种皮,外种皮细胞纵向延长,中种皮为7-9层切向延长的薄壁细胞,内种皮为石细胞型：合点区无合点室,内种皮在此出现缺口,缺口为整体轮廓呈喇叭形的近等径薄壁细胞群填充;珠孔区无珠孔领与孔盖的分化：外胚乳缺,内胚乳发达。蝎尾蕉属的Heliconia faranmansis?D6肿游藜僦制ぃ制の薹只墒闾寤闲∏揖断蜓映げ⑴帕形だ缸吹谋”谙赴钩桑褐榭锥酥制は蛲庋由欤纬衫嗨平浦肿拥闹指纷唇峁梗何蘅赘怯胫榭琢斓哂杏晒ば纬傻挠不牵缓系闱肼萌私断嗨疲煌馀呷樵?-4层细胞,细胞壁波浪形弯曲,内胚乳发达。综合作者对兰花蕉(Orchidanha chinensis)和前人对芭蕉群的种子解剖学研究结果,初步总结了芭蕉群种子解剖学特征及其进化式样,讨论了姜目芭蕉群四科种子解剖学特征的系统分类学意义。     

Weakening of muskmelon perisperm envelope tissue 4

Muskmelon (Cucumis melo L.) embryos are enclosed in an envelopeof tissue consisting of a layer of endosperm and a multi-cell-layeredperisperm that the radicle must penetrate for germination tooccur. The force and energy required to penetrate the perispermenvelope tissue were measured using an Instron universal testingmachine at a crosshead speed of 5 mm min^–1 after 0, 10,15, 22, 23, and 25 h of imbibition at 25C. The cellular structureof perisperm envelope tissue surrounding the radicle was observedafter 10, 15, 20, 25, and 48 h of imbibition using scanningelectron microscopy. The force required to puncture 5-mm-long,micropylar seed pieces declined steadily from 1.65 N in driedseeds to 0.65 N after 21 h of imbibition. The penetration energydeclined from 3.0 N mm in dry seeds to 1.1 N mm at 21 h afterthe start of imbibition when the first seeds germinated. Theforce and energy required to penetrate germinated seed pieceswere 0.55 N and 0.9 N mm, respectively, so the net punctureforce and energy needed to rupture the micropylar region ofthe perisperm envelope was roughly 0.10 N and 0.2 N mm at radicleemergence, respectively. Instron measurements of penetrationforce and energy decreased dramatically at crosshead speedsless than the 5 mm min^–1. Crosshead speeds greater than5 mm min^–1 may overestimate the pressure needed to ruptureperisperm and endosperm tissues. Intracellular cracks were firstobserved in SEM images 15 h after the start of imbibition, andafter 20 h cracking was apparent throughout the micropylar regionof the perisperm envelope. The perisperm envelope ruptured inone of two ways, coincident with radicle emergence. In approximately85% of muskmelon seeds, a large crack formed in the perispermenvelope adjacent to the radicle, while in roughly 15 % a circulararea of the perisperm envelope detached during radicle emergence.In dead seeds, the penetration force remained constant from10–24 h after the start of imbibition, and there wereno visible signs of tissue degradation. Cellular degradationand weakening of the perisperm envelope tissue precedes radicleemergence in muskmelon seeds. Key words: Seed, Instron, turgor, cell wall, electron microscopy, Cucumis melo     

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.     

Water Relations of Seed Development and Germination in Muskmelon (Cucumis melo L.) : IV. Characteristics of the Perisperm during Seed Development

We previously reported that an apparent water potential disequilibrium is maintained late in muskmelon (Cucumis melo L.) seed development between the embryo and the surrounding fruit tissue (mesocarp). To further investigate the basis of this phenomenon, the permeability characteristics of the tissues surrounding muskmelon embryos (the mucilaginous endocarp, the testa, a 2- to 4-cell-layered perisperm and a single cell layer of endosperm) were examined from 20 to 65 days after anthesis (DAA). Water passes readily through the perisperm envelope (endosperm + perisperm), testa, and endocarp at all stages of development. Electrolyte leakage (conductivity of imbibition solutions) of individual intact seeds, decoated seeds (testa removed), and embryos (testa and perisperm envelope removed) was measured during imbibition of freshly harvested seeds. The testa accounted for up to 80% of the total electrolyte leakage. Leakage from decoated seeds fell by 8- to 10-fold between 25 and 45 DAA. Presence of the perisperm envelope prior to 40 DAA had little effect on leakage, while in more mature seeds, it reduced leakage by 2- to 3-fold. In mature seeds, freezing, soaking in methanol, autoclaving, accelerated aging, and other treatments which killed the embryos had little effect on leakage of intact or decoated seeds, but caused osmotic swelling of the perisperm envelope due to the leakage of solutes from the embryo into the space between the embryo and perisperm. The semipermeability of the perisperm envelope of mature seeds did not depend upon cellular viability or lipid membrane integrity. After maximum seed dry weight is attained (35-40 DAA), the perisperm envelope prevents the diffusion of solutes, but not of water, between the embryo and the surrounding testa, endocarp, and mesocarp tissue.     

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。     

Seed development in Malpighiaceae species with an emphasis on the relationships between nutritive tissues

Malpighiaceae ovules have a well-developed nucellus; previous observations indicate that during seed development, the endosperm does not proliferate, thus, remaining scarce. This study aimed at identifying the nutritive tissues during seed development in Malpighiaceae, focusing especially on the endosperm. We analysed the seed development of Janusia mediterranea, J. occhionii, Mascagnia cordifolia, and Tetrapterys chamaecerasifolia, which were collected and processed by traditional methods for light microscopy. Ovules are subcampylotropous, crassinucellate and unitegmic in Janusia and bitegmic in M. cordifolia and T. chamaecerasifolia. The nucellus is well developed and protrudes through the micropyle, touching the funicular obturator. During development, a pachychalaza is formed, and the integuments coalesce in bitegmic species. Through a series of nucellar cell divisions, the perisperm is formed. In Janusia species, the endosperm is not produced. In M. cordifolia and T. chamaecerasifolia, the endosperm is nuclear, but it is scarce and ephemeral. The mature seed is exalbuminous, and the perisperm is consumed, and thus, the mature embryo is total. The absence of endosperm in Janusia is newly observed for the family and indicates functional transfer for the abundant perisperm.     

SEED MORPHOLOGY OF MONOTROPA UNIFLORA L. (ERICACEAE)

Mature seeds of Monotropa uniflora L., an achlorophyllous mycotrophic perennial, underwent imbibition and were processed for study using modern histological and histochemical techniques. The seeds ranged from 0.6-0.8 mm in length and 0.12-0.15 mm in width and exhibited integumentary winglike structures at either end. As in other members of the Ericaceae, seeds are unitegmic, tenuinucellate, albuminous, and form both micropylar and chalazal endosperm haustoria. A two-celled reduced embryo was observed in all seeds, except one where the embryo was three-celled. Protein granules in the thick walled endosperm were found to contain an aniline blue-fluorescent material that may be a calloselike carbohydrate. The limited amount of seed food reserves and the retardation of embryo differentiation may reflect specialized germination requirements.     

Ricinosomes provide an early indicator of suspensor and endosperm cells destined to die during late seed development in quinoa (Chenopodium quinoa)

Background and Aims

In mature quinoa (Chenopodium quinoa) seeds, the lasting endosperm forms a micropylar cone covering the radicle. The suspensor cells lie within the centre of the cone. During the final stage of seed development, the cells of the lasting endosperm accumulate protein and lipids while the rest are crushed and disintegrated. Both the suspensor and endosperm die progressively from the innermost layers surrounding the embryo and extending towards the nucellar tissue. Ricinosomes are endoplasmic reticulum-derived organelles that accumulate both the pro-form and the mature form of cysteine endopeptidase (Cys-EP), first identified in castor bean (Ricinus communis) endosperm during germination. This study sought to identify associations between the presence of ricinosomes and programmed cell death (PCD) hallmarks in suspensor and endosperm cells predestined to die during quinoa seed development.

Methods

A structural study using light microscopy and transmission electron microscopy was performed. To detect the presence of Cys-EP, both western blot and in situ immunolocalization assays were carried out using anti-R. communis Cys-EP antibody. A TUNEL assay was used to determine DNA fragmentation.

Results and Conclusions

Except for the one or two cell layers that constitute the lasting endosperm in the mature seed, ricinosomes were found in suspensor and endosperm cells. These cells were also the site of morphological abnormalities, including misshapen and fragmented nuclei, vesiculation of the cytosol, vacuole collapse and cell wall disorganization. It is proposed that, in suspensor and endosperm cells, the early detection of Cys-EP in ricinosomes predicts the occurrence of PCD during late seed development.