植物根中质外体屏障结构和生理功能研究进展

综述了近10年来植物根中质外体屏障结构和功能的研究进展。质外体屏障指根中内、外皮层初生壁的凯氏带,或次生壁栓质化和木质化,以及植物体表角质层组成的保护组织,能隔绝水、离子和氧气不能自由进出植物体的屏障结构,具有保护植物体的生理功能。根中凯氏带的分子发育机理研究表明根内皮层类似哺乳动物上皮组织的保护作用。植物根中质外体保证内部各种生理代谢在稳定的内部环境中进行,是植物适应各种逆境的重要屏障结构。根中质外体屏障在植物适应干旱、洪涝灾害、离子胁迫和病虫害的侵袭等方面具有重要作用,在探索适应并修复极端生态环境的植物资源中有广阔的应用前景。     

养分在植物体内循环的奥秘

植物的生长和发育离不开养分,其中所需的矿质营养主要来自根系的吸收。根系在供应植株地上部养分的同时,它还受到地上部养分需求的调节。因此植物根系和地上部之间保持着密切的联系,这种联系就是通过植物体内的养分循环来实现。植物体内的养分循环是指根系吸收的矿质营养,经木质部运输到地上部,其中一部分养分又经韧皮部返回根系的过程,即矿质营养经历了一个完整的循环过程:根系→木质部→地上部→韧皮部→根系。上述由地上部返回到根中的养分不能被根系完全利用,其中一部分又可经木质部再次运到地上部分,这一过程称为养分的再循环。根据矿…     

多细胞生物干细胞不对称分裂的内在调节机制研究进展

多细胞生物的发育是从一个受精卵分化成多种类型细胞的过程。细胞多样性形成的基础是不等分裂,不等分裂是干细胞自我更新和自我维持的关键。干细胞不等分裂有细胞内和细胞外两种调节机制。果蝇神经干细胞增殖和分化、植物胚胎发育、表皮气孔形成及根内皮层的分化,是研究不等细胞分裂调节机制最多的发育背景。本综述介绍了果蝇神经干细胞和植物胚胎发育早期、表皮气孔发生及根皮层内皮层中细胞不等分裂内在调节机制的研究进展。     

菰适应湿地环境的解剖和屏障结构特征研究

菰(Zizania latifolia)是一种多年生挺水植物,为了探讨该植物根、茎和叶的解剖结构、组织化学及其质外体屏障的通透性生理。该文利用光学显微镜和荧光显微镜,对菰的根、茎、叶进行了解剖学和组织化学研究。结果表明:(1)菰不定根解剖结构由外而内分别为表皮、外皮层、单层细胞的厚壁机械组织层、皮层、内皮层和维管柱;茎结构由外而内分别为角质层、表皮、周缘厚壁机械组织层、皮层、具维管束的厚壁组织层和髓腔。叶鞘具有表皮和具维管束皮层,叶片具有表皮,叶肉和维管束。(2)不定根具有位于内侧的内皮层及其邻近栓质化细胞和外侧的外皮层组成的屏障结构;茎具内侧厚壁机械组织层,外侧的角质层和周缘厚壁机械组织层组成的屏障结构,屏障结构的细胞壁具凯氏带、木栓质和木质素沉积的组织化学特点,叶表面具有角质层。(3)菰通气组织包括根中通气组织,茎、叶皮层的通气组织和髓腔。(4)菰的屏障结构和解剖结构是其适应湿地环境的重要特征,但其茎周缘厚壁层和厚壁组织层较薄。由此推测,菰适应湿地环境,但在旱生环境中分布有一定的局限性。     

女性生殖道微生态屏障

随着感染微生态学的发展,尤其是宿主-微生物相互作用的研究深入,越来越多的研究发现,新的微生态屏障在维持宿主健康方面发挥着重要作用。微生态屏障包括微生物屏障,机械屏障,免疫屏障和化学屏障四个部分。其中机械屏障是微生态屏障的基础,微生物屏障是微生态屏障的关键,免疫屏障是微生态屏障的核心,化学屏障是微生态屏障的有机组成部分。微生态屏障可维持宿主生境的微生态平衡,防止病原微生物的入侵和感染的发生。     

植物内皮层的发育与功能

在植物根的内皮层(某些茎的初生构造中也有)是皮层的最内层细胞。在结构和功能上,内皮层既不同于其内方的中柱鞘,也不同于外方的皮层薄壁组织。它的分化有着特殊的生理意义。植物组织发育的研究表明,内皮层来源于基本分生组织。但是,不同类群的植物,其内皮层的发育状况也不同。根中的内皮层常较为典型,它的发育一般终止于三种状态。第一种状态常见于大多数双子叶植物的幼根中,在幼小的内皮层细胞的径向壁和上、下端壁上,有一条初     

植物根系质外体屏障研究进展

根是植物吸收水分和矿质营养以维持生命活动的重要器官。根系的构型和超微结构具有物种特异性, 对水分和矿质营养的吸收有不同程度的影响。其中, 内、外皮层的木栓层和凯氏带是2种重要的质外体屏障, 可非定向地阻断水分和离子运输, 在植物生长发育及响应逆境胁迫中发挥重要作用。尽管如此, 植物根系质外体屏障的结构、化学组成、生理功能、生物合成及其调控仅在模式植物拟南芥(Arabidopsis thaliana)中被广泛研究。近年来, 关于作物大麦(Hordeum vulgare)、水稻(Oryza sativa)以及部分牧草的根系质外体屏障研究报道逐渐增多。该文系统比较了拟南芥、大麦、水稻以及部分牧草根系质外体屏障的异同, 提出今后的研究方向, 以期为深入探索禾本科作物和牧草根系质外体屏障在生长发育和逆境适应中的作用奠定理论基础, 并为作物和牧草育种工作提供新思路。     

喀斯特土壤条件下丛枝菌根真菌侵染对任豆幼苗生物量分配和根系结构特征的影响

喀斯特地区土壤瘠薄,植被恢复困难,根系对幼苗生长发育起重要作用。丛枝菌根真菌(AMF)侵染可促进喀斯特植物干物质积累及提升抗逆能力,但AMF对喀斯特地区主要恢复树种根系侵染机制及影响的研究尚不够深入。本研究以喀斯特地区典型造林树种任豆(Zenia insignis)幼苗为试验材料,利用不同养分条件的喀斯特原生土壤开展盆栽试验,以摩西球囊霉(Funneliformis mosseae,Fm)、根内球囊霉(Rhizophagus intraradices,Ri)和2种菌根真菌混合菌剂(mixture inoculation,Mi)进行接种,分析不同养分土壤条件下菌根真菌对任豆幼苗生物量分配和根系结构特征影响。结果表明:贫瘠喀斯特土壤条件下Fm处理根系侵染率最高,地上部生物量、根系生物量、总生物量、根尖数、分叉数、交叉数和投影面积较CK提高2.50、5.60、3.67、3.03、3.78、3.66和3.56倍;Ri处理对地上部生物量、根系生物量、总生物量、分叉数无显著影响,根尖数、交叉数、投影面积较对照降低46.59%、50.00%、38.60%;Mi处理地上部生物量、根系生物量、总生物量、分叉数、交叉数和投影面积较CK提高3.02、3.47、3.14、1.64、1.60和1.70倍,对根尖数无显著影响。较高养分喀斯特土壤条件下Fm处理根系生物量、投影面积较CK提高2.38、1.51倍,对地上部生物量、总生物量、分叉数和交叉数无显著影响,根尖数较CK降低22.82%;Ri处理根系生物量、总生物量、根尖数、交叉数和投影面积较CK提高2.29、1.37、1.31、1.29和1.66倍,对地上部生物量、根系分叉数无显著影响;Mi处理地上部生物量、总生物量、根尖数和投影面积较CK提高1.44、1.46、1.25和1.40倍,对根系生物量、分叉数和交叉数无显著影响。养分较低条件下,Fm侵染有利于任豆生物量提高和吸收性根系分化,有助于根系觅食养分和水分,促进植物生长;养分较高条件下,Ri促进根系结构特征改变,Fm促生效应减弱。低钙高磷养分条件下Ri表现出较强的促生作用;Fm生态适应性强,在促进植物根系获取土壤养分、水分及土壤空间拓展方面具有显著优势,可作为菌根真菌促进喀斯特地区植被恢复的优势菌种。     

数种植物生物电流日周期变化观测初报

植物对养分吸收的方式分为:主动吸收、被动吸收和胞饮吸收三种^[1]。在植物体内,几乎可发现所有存在于土壤中的各种元素,这充分说明了被动吸收对植物矿质养分吸收的重要性。在矿质养分被动吸收过程中,溶于水中的离子对,以水为载体进入植物体内。土壤化学称这股荷电的离子为离子流;对植物而言,进入植物体内的这股离子流即营养流^[2]。荷电的离子流源源不断地通过根系,进入植物体内,并定向地运往地上部,这个过程表现为可检测出的植物生物电流。由此可见,植物矿质养分的被动吸收,不仅受到蒸腾作用、光合作用等耗水代谢的影响,而且还受到电磁场、太阳辐射能和月球引力作用--潮汐等大地物理诸因子的影响^[4]。因此,研究植物体生物电的变化规律,可在一定程度上较综合地反映植物对矿质养分的总体吸收情况。为此,本试验拟对数种植物生物电流日周期变化进行初步探讨。     

SCFAs对肠道免疫调控的研究进展

肠道微生物群与宿主是共生关系,二者共同进化。短链脂肪酸(short-chain fatty acids, SCFAs)通过保护肠上皮屏障的完整性来维持肠道内环境平衡,并通过影响肠道免疫细胞的分化调节免疫系统。作为肠道微生物群发酵膳食纤维产生的一类重要代谢物,SCFAs通过抑制组蛋白脱乙酰酶或激活G蛋白偶联受体调节肠道免疫细胞功能与分化,在宿主的健康和免疫介导的疾病中发挥至关重要的作用。该文从SCFAs的来源、运输和信号转导,以及SCFAs对免疫细胞、免疫屏障及肠道疾病的影响等六个方面展开综述,并重点介绍了SCFAs对免疫细胞的作用。     

Comparative investigations on the differentiation of the endodermis and the development of the Hartig net in mycorrhizae of Picea abies and Larix decidua

Summary The differentiation of the endodermis of mycorrhizal roots of Picea abies and Larix decidua was investigated by means of light and transmission electron microscopy and with fluorescence techniques. The initiation and differentiation of the Hartig net were recorded. Differences between the two tree species were found, as were differences between the two tree species and angiosperms. The Casparian band developed immediately after the origin of endodermal cells from the meristem in mycorrhizae of both tree species. In L. decidua only the primary endodermis was present in most mycorrhizal laterals. The secondary structure of the endodermis was restricted to main roots and proximal parts of larch mycorrhizae. In P. abies mycorrhizae, however, the secondary stage of the endodermis developed soon after the primary endodermis and was characterized by regular alternation of short, active passage cells and elongated, rapidly degenerating cells, the inner surface of which was covered by a thick suberin layer. Hartig net development started in P. abies short roots only after the differentiation of endodermis into the secondary stage, whereas in L. decidua, the Hartig net was already initiated at the primary endodermal stage. Differences were specific for tree species.     

Chemical composition and ultrastructure of broad bean (<Emphasis Type="Italic">Vicia faba</Emphasis> L.) nodule endodermis in comparison to the root endodermis

Ultrastructure and development of apoplastic barriers within indeterminate root nodules formed by Vicia faba L. were examined by light and electron microscopy. The nodule outer cortex is separated from the inner cortex by a heavily suberized nodule endodermis, which matures in submeristematic regions and possesses suberin lamellae. Unsuberized passage cells are present near vascular strands, which are surrounded by a vascular endodermis attached on the inner side of the nodule endodermal cell walls. The vascular endodermis appears immediately below the meristematic apex in developmental state I (Casparian bands), gradually develops suberin lamellae, and attains developmental state II at the base of the nodule. For chemical analysis apoplastic barrier tissues were dissected after enzymatic digestion of non-impregnated tissues. Root epidermal and endodermal cell walls as well as nodule outer cortex could be isolated as pure fractions; nodule endodermal cell walls could not be separated from vascular endodermal cell walls and enclosed xylem vessels. Gas chromatography-flame ionization detection and gas chromatography-mass spectrometry were applied for quantitative and qualitative analysis of suberin and lignin in isolated cell walls of these tissues. The suberin content of isolated endodermal cell walls of nodules was approximately twice that of the root endodermal cell walls. The suberin content of the nodule outer cortex and root epidermal cell walls was less than one-tenth of that of the nodule endodermal cell wall. Substantial amounts of lignin could only be found in the nodule endodermal cell wall fraction. Organic solvent extracts of the isolated tissues revealed long-chain aliphatic acids, steroids, and triterpenoid structures of the lupeol type. Surprisingly, extract from the outer cortex consisted of 89% triterpenoids whereas extracts from all other cell wall isolates contained not more than 16% total triterpenoids. The results of ultrastructural and chemical composition are in good correspondence and underline the important role of the examined tissues as apoplastic barriers.     

Cluster root development in Grevillea robusta (Proteaceae). II. The development of the endodermis in a determinate root and in an indeterminate, lateral root

Light, fluorescence and electron microscopy were employed to follow the development of the endodermis in cluster roots and lateral roots of Grevillea robusta A. Cunn. ex R. Br. Endodermal cells had three different origins: rootlet endodermis arose from the rootlet meristem; endodermis covering the primordium shortly after initiation came from division of parental endodermis; cells at the junction between parent and rootlet endodermis developed from re-differentiated rootlet cortical cells. In the cluster root, the Casparian band formed in three ways, and was not initially present opposite the two sets of single xylem elements in the rootlet stele. A new clearing technique was developed that allowed visualization of xylem, suberized endodermis, Casparian band formation and phenolic compounds. In lateral roots, endodermal differentiation was asynchronous, but was related to position relative to protoxylem poles. However, the observed delay began before these poles had differentiated. At the tip of mature rootlets, which are determinate, the endodermis terminates in a 'dome' of cells, with the initial cell differentiating as an endodermal cell. Results are discussed in terms of determinate development in roots and the spatial and temporal contexts within which this development takes place.     

Focus Issue on Roots: Beyond the Barrier: Communication in the Root through the Endodermis

The root endodermis is characterized by the Casparian strip and by the suberin lamellae, two hydrophobic barriers that restrict the free diffusion of molecules between the inner cell layers of the root and the outer environment. The presence of these barriers and the position of the endodermis between the inner and outer parts of the root require that communication between these two domains acts through the endodermis. Recent work on hormone signaling, propagation of calcium waves, and plant-fungal symbiosis has provided evidence in support of the hypothesis that the endodermis acts as a signaling center. The endodermis is also a unique mechanical barrier to organogenesis, which must be overcome through chemical and mechanical cross talk between cell layers to allow for development of new lateral organs while maintaining its barrier functions. In this review, we discuss recent findings regarding these two important aspects of the endodermis.Soil contains water and dissolved nutrients needed for plant growth, but also holds pathogens and toxic compounds that can be detrimental to the plant. The root system, which is directly in contact with soil particles, can integrate environmental cues to adjust its development in order to optimize nutrient (Péret et al., 2011; Lynch, 2013) and water uptake (Cassab et al., 2013; Lynch, 2013; Bao et al., 2014) or avoid regions of high salinity (Galvan-Ampudia et al., 2013). Once anchored in the soil, roots must deal with the constraints of their local environment and develop specific barriers to balance uptake of nutrients, water, and interactions with symbionts with protection against detrimental biotic and abiotic factors.In young roots, these barriers are mainly formed by the deposition of hydrophobic polymers such as lignin and suberin within the primary cell wall of the endodermis, which separates the pericycle from the cortex (Fig. 1), and of the exodermis, which lies between the cortex and the epidermis (Nawrath et al., 2013). Although formation of an exodermis is species dependent, the endodermis is a distinguishing figure of extant vascular plants (Raven and Edwards, 2001). Within this layer, two barriers (i.e. the Casparian strip and the suberin lamellae) are sequentially deposited and regulate water and nutrient movements between the inner and outer parts of the root. In this review, we discuss how the presence of these two major endodermal barriers affects communication between the different cell layers of the root. We focus on recent articles highlighting the importance of the endodermis in this communication during various biological and developmental processes.Open in a separate windowFigure 1.Endodermal barriers affect radial movement of water and solutes through the root. A, At the root tip, to move from the soil to the outer tissues of the root and then into the stele, water and solute molecules can use either the apoplastic (black lines), symplastic (dotted lines), or transcellular (dashed lines) pathways. B, The deposition of the Casparian strip in the endodermis prevents the free apoplastic diffusion of molecules between the outer part and the inner part of the root forcing molecules to pass through the symplast of endodermal cells. C, The deposition of suberin lamellae prevents the uptake of molecules from the apoplast directly into the endodermis forcing molecules to enter the symplast from more outer tissue layers. Suberin deposition is also likely to prevent the backflow of water and ions out of the stele. Passage cells are unsuberized and may facilitate the uptake of water and nutrients in older parts of the root. Cor, Cortex; End, endodermis; Epi, epidermis; Peri, pericycle; Vasc, vasculature. Figure redrawn and modified from Geldner et al. (2013).     

The Casparian Strip of Clivia miniata Reg. Roots: Isolation,Fine Structure and Chemical Nature*

The root endodermis of Clivia miniata Reg. was successfully isolated using the cell wall degrading enzymes cellulase and pectinase. The enzymes did not depolymerize those regions of the primary cell walls of anticlinal endodermal root cells where the Casparian strips were located. Since the endodermis of C. miniata roots remained in its primary developmental state over the whole root length, endodermal isolates essentially represented Casparian strips. Thus, sufficient amounts of isolated Casparian strips could be obtained to allow further detailed investigations of the isolates by microscopic, histochemical and analytical methods. Scanning electron microscopy revealed the reticular structure of the Casparian strips completely surrounding the central cylinder of the roots. Whereas in younger parts of the root only the anticlinal cell walls of the endodermis remained intact in the isolates, in older parts of the root the periclinal walls also restricted enzymatic degradation due to the deposition of lignin. Extracts of the isolates with organic solvents did not reveal any wax-like substances which might have been deposited within the cell wall forming a transport barrier, as is the case with cutin and suberin. However, several histochemical and analytical methods (elemental analysis and FTIR spectroscopy) showed that the chemical nature of the Casparian strips of C. miniata roots can definitely be a lignified cell wall. These findings are in complete agreement with studies carried out at the beginning of this century on the chemical nature of the Casparian strips of several other plant species. The implications of these results concerning apoplasmatic transport of solutes and water across Casparian strips are discussed.     

The development of the endoder mis andPhi layer of apple roots

Summary Suberin lamellae and a tertiary cellulose wall in endodermal cells are deposited much closer to the tip of apple roots than of annual roots. Casparian strips and lignified thickenings differentiate in the anticlinal walls of all endodermal andphi layer cells respectively, 4–5 mm from the root tip. 16 mm from the root tip and only in the endodermis opposite the phloem poles, suberin lamellae are laid down on the inner surface of the cell walls, followed 35 mm from the root tip by an additional cellulosic layer. Coincidentally with this last development, the suberin and cellulose layers detach from the outer tangential walls and the cytoplasm fragments. 85 mm from the root tip the xylem pole endodermis (50% of the endodermis) develops similarly, but does not collapse. 100–150 mm from the root tip, the surface colour of the root changes from white to brown, a phellogen develops from the pericycle and sloughing of the cortex begins. A few secondary xylem elements are visible at this stage.Plasmodesmata traverse the suberin and cellulose layers of the endodermis, but their greater frequency in the outer tangential and radial walls of thephi layer when compared with the endodermis suggests that this layer may regulate the inflow of water and nutrients to the stele.     

Fourier transform infrared-spectroscopic characterisation of isolated endodermal cell walls from plant roots: chemical nature in relation to anatomical development

The chemical nature of enzymatically isolated endodermal cell walls from Cicer arietinum L., Clivia miniata Reg. and Iris germanica L. was studied by FTIR (Fourier transform infrared) spectroscopy. Observed frequencies were assigned to functional groups present in the cell wall and relative amounts of the biopolymers suberin and lignin, cell wall carbohydrates and proteins were determined. Infrared absorption spectra indicated structural characteristics for the three different developmental states of the isolated endodermal cell wall: primary endodermis with Casparian strips (state I), secondary endodermis with suberin lamellae (state II), and tertiary endodermis with U-shaped cell wall depositions (state III). The data obtained from this study are compared with previous results obtained by chemical degradation of isolated endodermal cell walls and subsequent determination of monomeric degradation products by gas chromatography and mass spectrometry. It is concluded that FTIR spectroscopy represents a direct and nondestructive method suitable for the rapid investigation of isolated plant cell walls. Furthermore, the observation that the suberin-assigned absorption bands disappeared after transesterification of the samples with BF₃-methanol confirmed that suberin is completely degraded by this treatment. Received: 20 February 1999 / Accepted: 25 May 1999