Root halotropism: Salinity effects on Bassia indica root

Abstract

Plant roots are responsible for the acquisition of nutrients and water from the soil and have an important role in plant response to soil stress conditions. The direction of root growth is gravitropic in general. Gravitropic responses have been widely studied; however, studies about other root tropisms are scarce. Soil salinity is a major environmental response factor for plants, sensed by the roots and affecting the whole plant. Our observations on root architecture of Kochia (Bassia indica) indicated that salinity may cue tropism of part of the roots toward increasing salt concentrations. We termed this phenomenon “positive halotropism”. It was observed that Kochia individuals in the field developed horizontal roots, originating from the main tap root, which was growing toward saline regions in the soil. Under controlled conditions in greenhouse experiments, Kochia plants were grown in pots with artificial soil salinity gradients, achieved by irrigation with saline and fresh water. It was shown that plants grown in low‐salt areas developed a major horizontal root toward the higher salt concentration region in the gradient. In regions of high salinity and in the absence of a salinity gradient, roots grew vertically without a major horizontal root. The novel finding of “positive halotropism” is discussed.     

Cadmium stress suppresses lateral root formation by interfering with the root clock

A biological clock activated by oscillating signals, known as root clock, has been linked to lateral root (LR) formation and is essential for regular LR spacing along the primary root. However, it remains unclear how this internal mechanism is influenced by environmental factors known to affect the LR pattern. Here, we report that excessive cadmium (Cd) inhibits LR formation by disrupting the lateral root cap (LRC)‐programmed cell death (PCD)‐regulated root clock. Cd restricts the frequency of the oscillating signal rather than its amplitude. This could be attributed to the inhibition on meristematic activity by Cd, which resulted in decreased LRC cell number and LRC‐PCD frequency. Genetic evidence further showed that LRC cell number is positively correlated with root resistance to Cd. Our study reveals root cap dynamics as a novel mechanism mediating root responses to Cd, providing insight into the signalling pathways of the root clock responding to environmental cues.     

The nutritional control of root development

Root development is remarkably sensitive to variations in the supply and distribution of inorganic nutrients in the soil. Here we review examples of the ways in which nutrients such as N, P, K and Fe can affect developmental processes such as root branching, root hair production, root diameter, root growth angle, nodulation and proteoid root formation. The nutrient supply can affect root development either directly, as a result of changes in the external concentration of the nutrient, or indirectly through changes in the internal nutrient status of the plant. The direct pathway results in developmental responses that are localized to the part of the root exposed to the nutrient supply; the indirect pathway produces systemic responses and seems to depend on long-distance signals arising in the shoot. We propose the term `trophomorphogenesis' to describe the changes in plant morphology that arise from variations in the availability or distribution of nutrients in the environment. We discuss what is currently known about the mechanisms of external and internal nutrient sensing, the possible nature of the long-distance signals and the role of hormones in the trophomorphogenic response.     

Intrinsic and environmental response pathways that regulate root system architecture

Root system development is an important agronomic trait. The right architecture in a given environment allows plants to survive periods of water of nutrient deficit, and compete effectively for resources. Root systems also provide an optimal system for studying developmental plasticity, a characteristic feature of plant growth. This review proposes a framework for describing the pathways regulating the development of complex structures such as root systems: intrinsic pathways determine the characteristic architecture of the root system in a given plant species, and define the limits for plasticity in that species. Response pathways co-ordinate environmental cues with development by modulating intrinsic pathways. The current literature describing the regulation of root system development is summarized here within this framework. Regulatory pathways are also organized based on their specific developmental effect in the root system. All the pathways affect lateral root formation, but some specifically target initiation of the lateral root, while others target the development and activation of the lateral root primordium, or the elongation of the lateral root. Finally, we discuss emerging approaches for understanding the regulation of root system architecture.     

The root of ABA action in environmental stress response

The growth and development of plants are influenced by the integration of diverse endogenous and environmental signals. Acting as a mediator of extrinsic signals, the stress hormone, abscisic acid (ABA), has been shown to regulate many aspects of plant development in response to unfavourable environmental stresses, allowing the plant to cope and survive in adverse conditions, such as drought, low or high temperature, or high salinity. Here, we summarize recent evidence on the roles of ABA in environmental stress responses in the Arabidopsis root; and on how ABA crosstalks with other phytohormones to modulate root development and growth in Arabidopsis. We also review literature findings showing that, in response to environmental stresses, ABA affects the root system architecture in other plant species, such as rice.     

In-situ root extent measurements by electrical capacitance methods

A conceptual model is presented that provides a rational basis for using plant root capacitance as an in-situ measurement for assessing plant root development. This method is based on measuring the electricla capacitance of an equivalent parallel resistance-capacitance circuit formed by the interface between soil-water and the plant root surface. Nutrient solution studies using tomato (Lycopersicon esculentum Mill.) showed a good correlation between plant root capacitance and root mass. Stage of development studies showed plant root capacitance measurements capable of detecting root development rate and suggested the method to be sensitive to root function. Soil water content was shown to have a significant effect on plant root capacitance measurement. The possibility of using this technique to assess relative root function is discussed. Positioning of the plant shoot electrode was shown to also have a significant effect on measurement of plant root capacitance, demonstrating the need for using consistent measurement techniques. The electrical capacitance method shows considerable promise. More research is needed before it can be used routinely.     

Genetic control of root exudation

The literature on genetics of root exudation and on genotypic differences in qualitative and quantitative composition of root exudates in crop and native plant species was critically assessed. Differences in exudation have been reported for genotypes that differ in tolerance to nutrient deficiencies, ion toxicities, and pathogen attack. The exudation profile of a limited number of genotypes (frequently only two genotypes with the contrasting response to the environmental stress) have been reported to date. Little is known about the variability in larger samples of the germplasm or about actual genetics behind differential qualitative and quantitative composition of root exudates. Changing the exudation profile of a given genotype may be achieved by manipulating the biosynthetic capacity and by increasing the capacity of the plasma membrane to transport the specific compound out into the rhizosphere. Overexpression of the bacterial citrate synthase gene in the cytoplasm of tobacco plants resulted in exudation of large quantities of citrate into the rhizosphere and partial alleviation of the aluminium (Al) toxicity stress. A similar strategy of transforming plants with citrate synthase gene is being tried as a way of improving plant capacity to extract phosphorus (P) from soils with notoriously low P availability.More research into the genetic basis of qualitative and quantitative differences in root exudation is warranted. Understanding the genetic control of root exudation, followed by manipulation of qualitative and quantitative composition of root exudates, will result in better adaptation of plants to environmental conditions and a greater yield of crops.     

System analysis of microRNAs in the development and aluminium stress responses of the maize root system

MicroRNAs (miRNAs) are a class of regulatory small RNAs (sRNAs) that down‐regulate target genes through mRNA cleavage or translational inhibition. miRNA is known to play an important role in the root development and environmental responses in both the Arabidopsis and rice. However, little information is available to form a complete view of miRNAs in the development of the maize root system and Al stress responses in maize. Four sRNA libraries were generated and sequenced from the early developmental stage of primary roots (PRY), the later developmental stage of maize primary roots (PRO), seminal roots (SR) and crown roots (CR). Through integrative analysis, we identified 278 miRNAs (246 conserved and 32 novel ones) and found that the expression patterns of miRNAs differed dramatically in different maize roots. The potential targets of the identified conserved and novel miRNAs were also predicted. In addition, our data showed that CR is more resistant to Al stress compared with PR and SR, and the differentially expressed miRNAs are likely to play significant roles in different roots in response to environmental stress such as Al stress. Here, we demonstrate that the expression patterns of miRNAs are highly diversified in different maize roots. The differentially expressed miRNAs are correlated with both the development and environmental responses in the maize root. This study not only improves our knowledge about the roles of miRNAs in maize root development but also reveals the potential role of miRNAs in the environmental responses of different maize roots.     

不同生境下荒漠植物红砂-珍珠猪毛菜混生根系的垂直分布规律

植物种间相互作用及其对环境胁迫的响应一直是物种共存和生物多样性维持研究的一个热点, 从地下根系入手来探讨混生群落植物种间关系及其对环境胁迫响应的研究少见报道。该文以荒漠草原区(灵武)、典型荒漠区(张掖)和极端荒漠区(酒泉) 3个不同生境条件下单生与混生红砂(Reaumuria soongarica)和珍珠猪毛菜(Salsola passerina)为实验材料, 采用分层取样法对其垂直根系参数进行测定和分析, 探讨了两种植物根系分布对混生及荒漠环境梯度的响应。结果表明: 同一生境条件下, 混生红砂和珍珠猪毛菜比根长和比表面积均高于单生, 说明红砂、珍珠猪毛菜混生后其根系相互作用关系表现为互惠, 促进了植株对土壤养分和水分的吸收。不同生境条件下, 同一生长方式的红砂根系分布深度均大于珍珠猪毛菜, 且根系消弱系数也普遍高于珍珠猪毛菜, 说明二者在不同生境条件下占据不同生态位, 红砂表现为深根性, 根系位于土壤深层, 珍珠猪毛菜表现为浅根性, 根系分布于土壤浅层。随着荒漠环境胁迫增强, 单生和混生红砂与珍珠猪毛菜的比根长和比表面积均呈现出极端荒漠区>典型荒漠区>草原荒漠区的规律, 且生境越干旱, 混生群落根系分离越明显; 单生与混生红砂根系消弱系数也逐渐增大, 在极端干旱区达到最大值, 珍珠猪毛菜变化不大, 表明红砂-珍珠猪毛菜混生群落根系生态位分离随荒漠环境胁迫增强而加大, 验证了环境胁迫梯度假说。可见“地上聚生, 地下分离”的混生方式可能是红砂-珍珠猪毛菜混生群落适应干旱胁迫环境的生长策略。     

类受体激酶调控根生长发育的研究进展

类受体激酶是一类具有激酶活性的单次跨膜受体,通过接收和传递胞外信号调控细胞的生理反应,参与植物生长发育过程。植物根在生长发育过程中受到大量的外部刺激和内源性发育信号的影响,植物必须通过整合这些信号并转化为细胞反应,才能适应不断变化的环境条件;植物类受体激酶作为细胞膜上的信息监测者,通过与外源和内源信号的通讯调控根的生长发育。该文对近年来国内外有关类受体激酶的结构、分类及其作用机制,特别是植物类受体激酶在根发育信号转导途径中的功能和作用等方面的研究进展进行综述,为进一步揭示植物类受体激酶在根生长发育中的功能及其作用机制提供参考。     

Cell-type action specificity of auxin on Arabidopsis root growth

The plant hormone auxin plays a critical role in root growth and development; however, the contributions or specific roles of cell-type auxin signals in root growth and development are not well understood. Here, we mapped tissue and cell types that are important for auxin-mediated root growth and development by manipulating the local response and synthesis of auxin. Repressing auxin signaling in the epidermis, cortex, endodermis, pericycle or stele strongly inhibited root growth, with the largest effect observed in the endodermis. Enhancing auxin signaling in the epidermis, cortex, endodermis, pericycle or stele also caused reduced root growth, albeit to a lesser extent. Moreover, we established that root growth was inhibited by enhancement of auxin synthesis in specific cell types of the epidermis, cortex and endodermis, whereas increased auxin synthesis in the pericycle and stele had only minor effects on root growth. Our study thus establishes an association between cellular identity and cell type-specific auxin signaling that guides root growth and development.     

Involvement of auxin pathways in modulating root architecture during beneficial plant–microorganism interactions

A wide variety of microorganisms known to produce auxin and auxin precursors form beneficial relationships with plants and alter host root development. Moreover, other signals produced by microorganisms affect auxin pathways in host plants. However, the precise role of auxin and auxin‐signalling pathways in modulating plant–microbe interactions is unknown. Dissecting out the auxin synthesis, transport and signalling pathways resulting in the characteristic molecular, physiological and developmental response in plants will further illuminate upon how these intriguing inter‐species interactions of environmental, ecological and economic significance occur. The present review seeks to survey and summarize the scattered evidence in support of known host root modifications brought about by beneficial microorganisms and implicate the role of auxin synthesis, transport and signal transduction in modulating beneficial effects in plants. Finally, through a synthesis of the current body of work, we present outstanding challenges and potential future research directions on studies related to auxin signalling in plant–microbe interactions.     

植物根压研究进展

根压是植物根部产生的一种静水压力,广泛存在于多种植物中。在蒸腾作用很弱的情况下,根压不但可驱动水分从根部流向冠层叶片,缓解因白天强烈蒸腾而导致的水分亏缺,而且在木质部导管栓塞修复方面具有重要作用。虽然国内外学者对根压的产生已有一些解释,普遍接受的观点有渗透理论、代谢理论和水分向上共同运输假说等,但根压产生的机制至今仍是科学家争议的焦点之一。根压的测定方法虽有直接和间接测定、损伤和无损伤测定之分,但较为先进的根压测定技术仍需进一步改善和提升。受水通道蛋白、遗传因素、生境等因素的影响,根压的大小存在差异,即使是较低的根压也会影响农作物生长。在促进转运蛋白质、酶、氨基酸、激素及钙元素等在农作物木质部和韧皮部之间流通方面,适当大小的根压发挥重要作用,且有助于提高农作物产量。因此,加深对植物根压的认识和理解具有重要的生物学意义。该文从根压的定义和产生机制、具有根压的植物类群、根压的测定方法和大小、影响根压的主要因素及根压在植物科学研究领域的意义和影响等多个方面分别进行了归纳总结,并结合当前研究热点和研究成果,针对植物根压研究过程中遇到的问题和后续研究趋势及方向进行了展望。     

不同抗旱性花生品种的根系形态发育及其对干旱胁迫的响应

为明确不同抗旱性花生品种的根系形态发育特征,探讨其根系形态发育特征对不同土壤水分状况的响应机制,在防雨棚旱池内进行土柱栽培试验,研究抗旱型品种“花育22号”、“唐科8号”和干旱敏感型品种“花育23号”3个不同抗旱性花生品种根系形态发育特征及其对干旱胁迫的响应.结果表明:抗旱型品种根系较发达,具有较大的根系生物量、总根长、总根系表面积.干旱胁迫使抗旱型品种根系总表面积和体积增加,而干旱敏感型品种则相反.干旱胁迫显著增加抗旱型品种“花育22号”20 cm以下土层内根长密度分布比例及根系表面积和体积,但“唐科8号”相应根系性状仅在20-40 cm土层内增加;干旱胁迫使干旱敏感型品种“花育23号”40 cm以下土层内各根系性状升高,但未达显著水平且其深层土壤内各根系性状增加幅度小于“花育22号”.花生根系总长、总表面积及0-20 cm土层内根系性状与产量间呈显著或极显著正相关.土壤水分亏缺条件下,花生主要通过增加深层土壤内根长、根系表面积和体积等形态特性,优化空间分布构型,以调节植株对水分的利用.     

Modification of root architecture in woody plants is possible for the presence of two different mechanisms of lateral root production: The effect of slope in Spartium junceum L. seedlings

Abstract

This paper investigates the modification of root architecture of Spartium junceum L. seedlings grown in slope condition. It is reported that 50% of the total number of lateral roots are concentrated in few centimetres of the taproot near the collar. The anatomical analysis of transverse sections along the taproot axis reveals that this taproot zone is characterised by two types of lateral roots: one with a trace extending to the centre of the vascular cylinder by following the path of a medullar ray; one with a trace which ends in the vascular cambium. The first type may be lateral roots originated from the taproot primary structure; the second type seems to be lateral roots developing later when a secondary structure has completely substituted the primary structure. The emission of this second type of lateral roots seems to be strongly controlled by environmental conditions with considerable consequences upon the overall root architecture. In the example reported in this paper, young plants growing under mechanical stress due to a slope develop asymmetric root architecture with lateral roots elongating in two prevalent directions: up-slope or down-slope. This asymmetric architecture is produced in the zone of the taproot where a secondary structure is present and represents the plant response to the need of increasing its anchorage strength.     

Variations of root traits in three Xizang grassland communities along a precipitation gradient

根系功能属性及其变异性能够介导物种共存及环境适应策略, 但强烈的环境约束作用能够引起不同物种间根系属性的趋同性。为了研究西藏高寒草原群落中植物根系属性变异规律, 并阐明不同物种资源获取和适应策略的多样性, 该文对西藏高寒草原不同的环境梯度进行了研究。作者自东向西沿着降水梯度在那曲、班戈和尼玛3个自然草原群落进行群落调查, 并采集了共计22种植物。测定了每种植物的一级根直径、一级侧根长度和根系分支强度3个关键根系属性。结果表明: 在西藏高寒草原群落中, 不同物种根系直径普遍较小, 且种间变异非常小(22.76%), 其中86%的物种一级根直径集中在0.073 mm到0.094 mm之间; 相较于直径较粗的物种, 直径越细的物种分支强度越高, 侧根越短。在群落尺度上, 植物主要通过增加根系直径、侧根长度, 降低分支强度的方式来适应水分的减少; 而在物种尺度上, 植物适应水分变化的策略则呈现多样性。     

西藏高寒草原群落植物根系属性在降水梯度下的变异格局

根系功能属性及其变异性能够介导物种共存及环境适应策略, 但强烈的环境约束作用能够引起不同物种间根系属性的趋同性。为了研究西藏高寒草原群落中植物根系属性变异规律, 并阐明不同物种资源获取和适应策略的多样性, 该文对西藏高寒草原不同的环境梯度进行了研究。作者自东向西沿着降水梯度在那曲、班戈和尼玛3个自然草原群落进行群落调查, 并采集了共计22种植物。测定了每种植物的一级根直径、一级侧根长度和根系分支强度3个关键根系属性。结果表明: 在西藏高寒草原群落中, 不同物种根系直径普遍较小, 且种间变异非常小(22.76%), 其中86%的物种一级根直径集中在0.073 mm到0.094 mm之间; 相较于直径较粗的物种, 直径越细的物种分支强度越高, 侧根越短。在群落尺度上, 植物主要通过增加根系直径、侧根长度, 降低分支强度的方式来适应水分的减少; 而在物种尺度上, 植物适应水分变化的策略则呈现多样性。