传递细胞——物质短途运输中起作用的特化细胞

传递细胞广泛存在于植物界的各种群。传递细胞的分化主要与器官的发育程度以及转运物质的供应有关。当植物某个部位所需的运输速率远高于溶质正常跨膜运输速率时,在此部位就可能有传递细胞。传递细胞最基本的特征是细胞壁向内突起生长（壁内突）并与质膜共同形成壁膜器。壁内突从形态上可划分为2种类型：网状内突和肋状内突。大多数传递细胞壁内突的发育在沿着溶质流动的方向表现出极性。传递细胞的胞质一般比周围薄壁组织细胞浓,胞内富含线粒体和内膜分泌系统细胞器如内质网、高尔基体、小囊泡等。传递细胞在物质的短途运输中起作用。玉米胚乳传递细胞可能还具有防御病原微生物进入胚乳和胚的功能。本文就传递细胞的种类和特性、结构和功能、形成机制和诱导因素,以及基因表达调控等方面的研究进展做介绍。     

矮生菜豆胚乳细胞形成时的超微结构研究

应用透射电镜对矮生菜豆的游离核胚乳、胚乳细胞及珠被绒毡层的超微结构进行了观察,揭示了胚乳细胞化过程中自由生长壁的形成方式。实验表明,在游离核胚乳中存在大量有被小泡,胚乳游离核的外层核膜发生明显外突并以出芽的方式产生有被小泡。初始胚乳细胞壁为自由生长壁,大量有被小泡融合于自由生长壁末端使之不断延伸并产生分枝。与珠被绒毡层相邻的胚乳细胞产生发达的壁内突,具有传递细胞特征;在这些细胞中出现大量的核糖体及粗糙内质网,表现现高度的合成活性。     

薏苡胚乳传递细胞的超微结构

传粉后１０ ｄ,薏苡（Ｃｏｉｘ ｌａｃｒｙｍ ａ－ｊｏｂｉＬ．）颖果基部胚乳最外层传递细胞具长而多的壁内突,第二层细胞的壁内突较第一层的短而少,均具瓣裂的细胞核、丰富的线粒体、粗糙内质网、核糖体、产生小泡的高尔基体及与壁内突质膜相连的、含深色物质的囊泡。线粒体分布于壁内突附近或其间。授粉后２５ ｄ,第一、二层细胞壁内突发达,几乎充满了细胞,但细胞器可见。第四层传递细胞具树枝状及网状的壁内突,大量线粒体、具质体膜的淀粉粒、脂体存在壁内突附近或壁内突的间隙内。高尔基体常见,仅见很少的片段内质网。第五层传递细胞具短的壁内突、较大的淀粉粒及许多小蛋白质体。两个时期的第一、二层细胞内均未观察到胞间连丝。授粉后２５ ｄ,第四层及以上的传递细胞的细胞壁和呈网状的壁内突均含有胞间连丝。还讨论了各种细胞器的作用及各层传递细胞的功能     

豇豆胚胎发育早期的胚柄及胚乳中的传递细胞

在豇豆早期的胚胎发育中,随着胚柄和胚乳在结构上的分化,可以看到在特定部位上形成传递细胞特有的壁内突。在原胚早期,只在胚周围和合点处的胚囊周界壁上存在壁内突。在原胚后期,胚柄的基部细胞的侧壁以及与母体相邻的端壁都发育壁内突。这个时期与珠心冠和内珠被相邻的胚乳细胞的外切向壁上也开始发生内突。胚柄在胚分化时期明显地分化为基部区域和颈部区域。壁内突只发生在基部区域的细胞中。在原胚后期,胚周围形成细胞胚乳的鞘。这部分的胚乳,在种于发育时期,细胞的数目和大小都只稍有增加。当胚乳细胞形成以后,细胞很快变为彼此分离,与珠被组织直接相邻的胚乳细胞的壁发生明显的内突。胚分化和胚轴延长时期,在液体胚乳的区域,在向珠柄那一面的周界壁上,出现特别发达的壁内突。     

根瘤形成过程中豆科植物根毛和根外层传递细胞的诱发

对7种豆科植物接种根瘤菌后根部的形态和内部结构进行了研究.结果表明:根瘤菌可诱发根瘤形成部位根段的根毛增生、形变和根外层传递细胞的发育.根外层传递细胞发生在根毛伸长形变时期,一直可持续到根瘤形成,传递细胞壁内突发育过程是先由根表皮细胞外切向壁一侧细胞质膜向细胞质内陷形成囊状壁傍体,次生细胞壁物质在初生壁上沉积并逐渐充满囊状体,最终形成传递细胞典型的壁内突结构.根瘤形成过程中根外层传递细胞的诱发与培养方式(水培、固培)没有直接关系.在不接菌的对照苗的根段内未发现壁内突结构,研究证明豆科植物根外层传递细胞的形成是由根瘤菌诱导所致.     

长豇豆胚和胚乳的发育及营养物质积累

长豇豆(Vigna sesquipedalis (L.)Fruwirth)开花前7—10小时传粉,开花后8—10小时完成双受精。合子期珠孔端及合点部位胚囊的周界壁有壁内突。胚发育属柳叶菜型。胚柄的基部细胞及基部区域外层细胞的外切向壁发生壁内突。成熟胚中胚柄宿存。开花后9—16天为子叶细胞中淀粉积累期,开花后12—18天为蛋白质积累期。胚乳发育为核型,珠孔端胚乳细胞化,合点端保持游离核状态。胚乳外层细胞为传递型细胞,珠孔端的胚乳细胞形成折叠细胞群,亦有壁内突。心形胚期胚乳开始退化解体,成熟胚期胚乳完全消失。     

芡实种子萌发期子叶传递细胞发育的初步研究

芡实种子萌发期,子叶吸收外胚乳中养分供萌发和幼苗发育,具有吸器的功能。在种子萌发过程中,子叶的部分表皮细胞发育为传递细胞。其壁内突的生长以外切向壁为多,形成壁内突的造壁物质主要由高尔基体合成,并由其溢出的囊泡运送的。     

饭包草种子萌发过程中养分运输和 消耗的细胞学过程初探

对饭包草大粒种子萌发过程中胚结构变化、养分运输和肖耗的细胞学过程进行了观察.结果显示:种子萌发时萌发孔盖张开,胚根、胚芽、胚轴、子叶鞘和子叶柄突破种皮生长,子叶片呈球状留在胚乳内吸收营养;透射电镜观察显示胚乳养分运输主要通过传递细胞运输、穿壁运输、胞间通道运输三种方式;细胞内养分消耗过程是细胞程序性死亡的过程,多膜小体...     

天竺葵花粉营养细胞的传递细胞型特征

天竺葵花粉营养细胞的传递细胞型特征罗玉英,胡适宜（北京大学生命科学学院,北京１００８７１）传递细胞型细胞在生殖系统中广泛存在,但这种细胞型具壁内突的特征,在花粉壁的许多研究中尚少报道,至今只在黑麦中有详细的描述。本文报道天竺葵花粉的营养细胞具传递细胞...     

花后低温条件下小麦颖果发育的显微结构观察北大核心CSCD

该研究以春性小麦品种‘扬麦15’和半冬性小麦品种‘烟农19’为实验材料,在小麦花后6~8 d进行低温处理,对花后不同发育阶段的小麦颖果进行取样,利用树脂切片技术观察了果皮、胚乳和养分运输组织的显微结构,以探明花后低温条件下小麦颖果发育的显微结构特征。结果表明:(1)花后低温延缓了两个小麦品种颖果早期和中期发育过程,推迟了‘扬麦15’灌浆期,缩短了‘烟农19’的灌浆期,最终降低了两小麦品种的粒重。(2)花后低温减缓了小麦颖果发育早期果皮细胞的凋亡以及中果皮细胞中淀粉体的降解速率。(3)花后低温明显降低了小麦胚乳发育早期和中期淀粉体和蛋白体的积累量,缩短了灌浆时期,降低了胚乳充实度。(4)花后低温条件下小麦颖果韧皮部发育缓慢,筛管分布范围缩小;同时胚乳传递细胞的发育变慢,细胞壁上壁内突数量明显减少。研究发现,花后低温使得两小麦品种颖果早期发育过程延缓,具体表现在果皮的凋亡推迟,贮藏物质积累量减少,养分运输组织变得不发达,胚乳充实度降低,单粒颖果干重下降;同时两小麦品种对低温的响应又存在差异,花后低温下春性小麦颖果灌浆期被延长,半冬性小麦灌浆期被缩短。     

Development of flange and reticulate wall ingrowths in maize (Zea mays L.) endosperm transfer cells

Maize (Zea mays L.) endosperm transfer cells are essential for kernel growth and development so they have a significant impact on grain yield. Although structural and ultrastructural studies have been published, little is known about the development of these cells, and prior to this study, there was a general consensus that they contain only flange ingrowths. We characterized the development of maize endosperm transfer cells by bright field microscopy, transmission electron microscopy, and confocal laser scanning microscopy. The most basal endosperm transfer cells (MBETC) have flange and reticulate ingrowths, whereas inner transfer cells only have flange ingrowths. Reticulate and flange ingrowths are mostly formed in different locations of the MBETC as early as 5 days after pollination, and they are distinguishable from each other at all stages of development. Ingrowth structure and ultrastructure and cellulose microfibril compaction and orientation patterns are discussed during transfer cell development. This study provides important insights into how both types of ingrowths are formed in maize endosperm transfer cells.     

Evidence for factors regulating transfer cell-specific expression in maize endosperm

In maize, a layer of basal endosperm cells adjacent to the pedicel is modified for a function in solute transfer. Three genes specifically expressed in this region, termed the basal endosperm transfer layer (BETL-2 to -4), were isolated by differential hybridization. BETL-2 to -4 are coordinately expressed in early and mid-term endosperm development, but are absent at later stages. BETL-2 to -4 coding sequences all predict small (<100 amino acids), secreted, cysteine-rich polypeptides which lack close relatives in current database accessions. BETL-3 and BETL-1 display some sequence similarities with each other and to plant defensins. BETL-2 to -4 promoter regions were isolated and compared, revealing the presence of a promoter-proximal microsatellite repeat as the most highly conserved sequence element in each sequence. Electrophoretic mobility shift assays (EMSA) showed that specific BETL-2 to -4 promoter fragments competed for binding to the same DNA-binding activity in nuclear extracts prepared from maize endosperm. Although BETL-2 to -4 are only expressed in basal endosperm cells, the DNA-binding activities detected were of two types: distal endosperm-specific, or present in both basal and distal endosperm extracts. On the basis of these findings, a model to account for the coordinate regulation of BETL genes in endosperm cells is proposed.     

Development and function of caryopsis transport tissues in maize,sorghum and wheat

Cereal caryopsis transport tissues are essential channels via which nutrients are transported into the embryo and endosperm. There are differences and similarities between caryopsis transport tissues of maize, sorghum and wheat. Vascular bundle, endosperm transfer cells, endosperm conducting cells and embryo surrounding region are common in maize, sorghum and wheat. Placentochalaza is special in maize and sorghum, while chalaza and nucellar projection transfer cells are special in wheat. There is an obvious apoplastic cavity between maternal and filial tissues in sorghum and wheat caryopses, but there is no obvious apoplastic cavity in maize caryopsis. Based on the latest research, the development and function of the three cereal caryopsis transport tissues are discussed and investigated in this paper.     

A Comparative Study on the Role of Cytokinins in Caryopsis Development in the Maize miniature1 Seed Mutant and Its Wild Type

We report here on a comparative developmental profile of plant hormone cytokinins in relation to cell size, cell number and endoreduplication in developing maize caryopsis of a cell wall invertase-deficient miniature1 ( mn1 ) seed mutant and its wild type, Mn1 , genotype. Both genotypes showed extremely high levels of total cytokinins during the very early stages of development, followed by a marked and genotype specific reduction. While the decrease of cytokinins in Mn1 was associated with their deactivation by 9-glucosylation, the absolute and the relative part of active cytokinin forms was higher in the mutant. During the exponential growth phase of endosperm between 6 d after pollination and 9 d after pollination, the mean cell doubling time, the absolute growth rate and the level of endoreduplication were similar in the two genotypes. However, the entire duration of growth was longer in Mn1 compared with mn1 , resulting in a significantly higher cell number in the Mn1 endosperm. These data correlate with the previously reported peak levels of the Mn1 -encoded cell wall invertase-2 (INCW2) at 12 d after pollination in the Mn1 endosperm. A model showing possible crosstalk among cytokinins, cell cycle and cell wall invertase as causal to increased cell number and sink strength of the Mn1 developing endosperm is discussed.