红树胎生现象及其对潮间带生境适应性研究进展

热带亚热带海岸潮间带的高盐、淹水、土壤缺氧和潮水冲击等不良环境因子导致了红树植物在形态、生理和生态方面的特异性 ,尤其是其独特的胎生现象。从红树植物胎生繁殖体在母体的发育过程和繁殖体落地后对潮间带生境的适应两个角度探讨了红树植物胎生现象的适应意义。红树植物的胎生现象是对潮间带盐胁迫、淹水、土壤缺氧和潮水冲击等综合生境的适应 ,而不是单独其中一种胁迫因子的适应。红树植物胎生繁殖体在母体的发育过程中通过能量积累、渗透调节、形态完善等过程为掉落以后适应潮间生境做准备 ,掉落的胎生繁殖体依靠从母体获得的能量和营养克服潮间带的盐胁迫、淹水、土壤缺氧和潮水冲击等多重胁迫。红树植物胎生繁殖体落地后的能量收支平衡是决定繁殖体定居成功与否的关键 ,而胎生现象的意义正是通过胎生繁殖体在母体发育过程中积累足够的能量和营养来适应潮间带生境。红树植物胎生繁殖体落地后苗木在潮间带滩涂的生长发育过程中存在一个能量“阈限”,能阈的高低决定于潮间带生境的恶劣程度 ,而突破能阈能力决定于胎生繁殖体所携带的能量与营养的数量。这为红树林湿地恢复提供了理论基础 ,但还需要更多的证据来支持该理论     

植物淹水胁迫的生理学机制研究进展

淹水胁迫引起弱光环境,使气体扩散受限,叶片细胞膜脂过氧化加剧,体内保护酶系统受损,叶绿素降解,丙二醛含量积累,光合速率下降。为了适应淹水环境,植物通过生理生化机制的调节来保证淹水条件下的生命活动。如细胞通过调节渗透物质的含量来保持渗透势的平衡;细胞内各种抗氧化酶活性增加,以清除自由基,避免或者减轻细胞受到伤害;改变代谢途径和激素调节以保持能量储备和低的代谢速率。本文综述了淹水胁迫对细胞膜系统及功能、植物光合作用、植物呼吸、激素、生理代谢、基因调控的影响和淹水结束后植物的生理生态学变化,介绍了植物适应淹水胁迫的机制,并指出植物耐淹响应的分子机理,环境因素对淹没植物的影响,森林淹水胁迫的定位观测是今后需要研究的方面。     

根际氧含量影响植物生长的生理生态机制研究进展

氧是植物生命活动所必需的营养因子,充足的氧供应才能满足植物正常生长发育的需求,但频发的洪涝灾害使根系淹水缺氧成为植物生长过程中经常遭受的非生物胁迫。因此,了解植物对根际氧含量变化的响应与适应机制对作物的生产及抗性品种的选育有重要意义。该文对根际氧环境对植物多方面的影响进行了综述,包括植物的需氧量、氧含量对根际环境的影响和氧含量对植物的影响等,并阐明植物在低氧下的伤害与适应机理和植物对低氧信号的感应与信号传导机制,最后总结了避免缺氧伤害的措施,同时还提出根际氧含量方面有待进一步探索的问题与研究方向,为后续的研究提供新思路。     

植物应对干旱胁迫的阶段性策略

干旱是影响植物生存、生长和分布的最重要的非生物胁迫之一,全球暖干化将加剧干旱胁迫.植物对干旱胁迫的响应和适应机制一直是
学术研究的热点领域.本文综述了植物应对干旱胁迫的生长和生理响应,在已有的研究结果基础上,提出了植物应对干旱胁迫的阶段性响应策略.从干旱开始到干旱致死,植物经历了干旱开始-轻度干旱-中度干旱-严重干旱-极端干旱5个阶段,分别对应着应激响应-主动适应-被动适应3种响应方式和适应机制.不同阶段中植物抗旱机制的核心任务不同.最后提出了研究植物阶段性响应策略需要解决的关键科学问题及研究方向.     

五种红树植物通气组织对人工非潮汐生境的响应

红树植物自然条件下生长于河口、海岸潮间带。受潮汐作用影响,红树植物在生理、形态、结构上对渍水环境产生了相应的适应机制。其中红树植物通气组织的发达程度与其耐淹水的能力具有很高的相关性,是衡量红树植物耐淹浸能力的重要依据。利用测定孔隙率和石蜡切片面积比两种方法揭示了华南地区5种红树植物优势种：白骨壤 (Avicennia marina)、红海榄 (Rhizophora stylosa)、木榄 (Bruguiera gymnorrhiza)、秋茄 (Kandelia candel)和桐花树 (Aegiceras corniculatum)在自然条件和人工生境下根通气组织的发育规律,并用石蜡切片研究了茎和叶的通气组织发育状况。结果表明：两种方法测得根的通气组织发育程度的结果相关性显著（P<0.05）。5种红树植物通气组织主要产生于根部,茎和叶发育较少,除了潮汐生境中白骨壤根的通气组织为根、茎、叶总和的48.16%、非潮汐生境中桐花树根为43.81%,其余树种根部通气组织占总体的50%以上。自然潮间带生境中,桐花树、木榄、白骨壤、秋茄、红海榄,通气组织分别为(14.98±3.34)%、(27.83±2.3)%、(29.64±3.17)%、(3009±4.12)%、(42.12±3.14)%,通气组织比例与其在潮间带上的分带性和演替序列较为吻合。非潮汐人工生境下,红海榄、木榄、秋茄、桐花树和白骨壤根部通气组织较自然生境下均有所增加,说明各树种对非潮汐淹浸条件具备一定的适应力。根据非潮汐生境下通气组织的比例可判定它们对恒定水位的适应能力依次为：桐花树>白骨壤>秋茄>木榄>红海榄。红树植物对非潮汐淹浸条件的适应有利于在沿海地区开展人工生境下红树林的栽培与推广应用,研究结果对提高栽培成活率,更大限度地发挥红树林的生态服务价值,具有重大的实践意义。     

红树植物水分关系研究进展

红树林生长于受潮汐影响的海滨特殊生境,具有重要的生态功能及应用价值。红树植物的水分利用特点一直是研究热点。由于受环境盐分影响,红树植物水势普遍较低;水分运输系统抗气穴化能力极强;水分利用保守,用水量处于同径级的热带陆生树木用水量的低值范围;表型可塑性大,可通过耐旱、耐盐和多样的水分管理策略适应潮间带环境;水分运输效率不低,能在环境条件适宜时进行高效的光合合成。本文通过大量的文献分析,综述了红树植物的水分关系特点、水分和盐分管理策略,对未来从多角度、结合新研究技术的红树植物水分关系研究进行了展望。     

红树植物水分关系研究进展

红树林生长于受潮汐影响的海滨特殊生境,具有重要的生态功能及应用价值。红树植物的水分利用特点一直是研究热点。由于受环境盐分影响,红树植物水势普遍较低;水分运输系统抗气穴化能力极强;水分利用保守,用水量处于同径级的热带陆生树木用水量的低值范围;表型可塑性大,可通过耐旱、耐盐和多样的水分管理策略适应潮间带环境;水分运输效率不低,能在环境条件适宜时进行高效的光合合成。本文通过大量的文献分析,综述了红树植物的水分关系特点、水分和盐分管理策略,对未来从多角度、结合新研究技术的红树植物水分关系研究进行了展望。     

红树植物胎生现象研究进展

植物胎生是指有性繁殖产生的后代在母体上直接萌发的现象, 在红树植物中最为常见。红树植物生长在热带亚热带海岸潮间带, 耐受高盐、高温、淹水缺氧和海浪冲击等复杂环境。胎生被认为是红树植物对这种特殊生境的重要适应方式。该文从形态发育、生理生化、分子水平、生态适应4个层次讨论红树植物胎生现象对复杂生境的适应性, 并指出现有研究存在的不足, 对将来的研究方向进行了展望。与非胎生胚胎发育相比, 红树植物胎生是一个遗传的程序, 在进化过程中形成了一些特殊的结构。植物激素对胎生发育起关键的调控作用, 繁殖体发育过程中, 其盐离子的种类与浓度的动态变化则是对海岸潮间带生境的重要适应特征。这种胎生繁殖体依靠在母体上完善的一系列功能性特征能更有效地适应落地后的滩涂环境。然而, 红树植物胎生发育过程的分子机理及调控机制还有待研究。理解胎生这一特殊适应性现象的本质及其进化过程将为红树林保护繁育、适应气候变化提供理论依据。     

植物涝渍胁迫应激机制研究进展

涝渍胁迫使植物从有氧呼吸转变为无氧呼吸,为了适应水淹环境,植物会根据自身特点启动应激机制。从植物形态、植物生理代谢、植物应激反应以及植物根际土壤微生物的变化阐述了植物对涝渍胁迫的应激机制,并对微生物生态响应机制与涝渍胁迫的关系方面的研究提出了展望,以期为植物抗性应激机制研究提供参考。     

扩展蛋白与植物抗逆性关系研究进展

扩展蛋白是一种细胞壁蛋白,可调节细胞壁的松弛和伸展。目前研究表明,扩展蛋白几乎参与调节植物生长发育的整个进程。扩展蛋白还与植物的多种抗性反应有关,在植物对干旱、高盐以及病虫害等生物胁迫和非生物胁迫响应方面起着重要的调节作用。干旱胁迫下扩展蛋白基因的表达与植物的抗旱性有一定的关系;植物的耐盐性受到扩展蛋白基因表达的影响;淹水促进植物的伸长生长与扩展蛋白的表达密切相关;扩展蛋白调节细胞壁松弛为植物抗病性研究提供了新的思路。     

Contrasting growth and adaptive responses of two oak species to flooding stress: role of non-symbiotic haemoglobin

Soil flooding is an environmental constraint that is increasingly important for forest ecosystems, affecting tree growth and regeneration. As a result, selection pressure will alter forest diversity and distribution by favouring tree species tolerant of soil oxygen deprivation. Sessile and pedunculate oaks are the most abundant oak species and they exhibit a strong differential tolerance to waterlogging. In order to gain some understanding of the mechanisms of tolerance of both species to hypoxia, we undertook the characterization of the physiological, morphological, cellular and molecular responses of both species to flooding stress. Our results indicate that pedunculate oak, the more tolerant species, succeeded in maintaining its growth, water status and photosynthetic activity at a higher level than sessile oak. Furthermore, pedunculate oak developed aerenchyma in its root cortex as well as adventitious roots. The later exhibited a strong accumulation of class1 non-symbiotic haemoglobin localized by in situ hybridization in the protoderm and in some cortical cells. In conclusion, the higher tolerance of pedunculate oak to flooding was associated with an enhanced capacity to maintain photosynthesis and water homeostasis, coupled with the development of adaptive features (aerenchyma, adventitious roots) and with a higher expression of non-symbiotic haemoglobin in the roots.     

Physiology and biochemistry of waterlogging tolerance in plants

Waterlogging is a serious problem, which affects crop growth and yield in low lying rainfed areas. The main cause of damage under waterlogging is oxygen deprivation, which affect nutrient and water uptake, so the plants show wilting even when surrounded by excess of water. Lack of oxygen shift the energy metabolism from aerobic mode to anaerobic mode. Plants adapted to waterlogged conditions, have mechanisms to cope with this stress such as aerenchyma formation, increased availability of soluble sugars, greater activity of glycolytic pathway and fermentation enzymes and involvement of antioxidant defence mechanism to cope with the post hypoxia/anoxia oxidative stress. Gaseous plant hormone ethylene plays an important role in modifying plant response to oxygen deficiency. It has been reported to induce genes of enzymes associated with aerenchyma formation, glycolysis and fermentation pathway. Besides, nonsymbiotic-haemoglobins and nitric oxide have also been suggested as an alternative to fermentation for maintaining lower redox potential (low NADH/NAD ratio), and thereby playing an important role in anaerobic stress tolerance and signaling.     

Photosynthetic and physiological responses of Kandelia candel L. Druce seedlings to duration of tidal immersion in artificial seawater

The present study demonstrates the influence of the duration of periodical waterlogging with artificial seawater on the photosynthetic and physiological responses of Kandelia candel L. Druce seedlings, the pre-dominant species of subtropical mangroves in China. Artificial tidal fluctuations applied here closely mimicked the twice daily tidal inundation which mangroves experience in the field. All the seedlings were immersed in artificial seawater during 70-day cultivation. Similar trends with increasing duration of immersion occurred in photosynthetic rate, transpiration rate, stomatal conductance and intercellular CO₂ concentration, where significant decreases occurred only in long time treatments of 10 or 12 h. Water used efficiency and chlorophyll contents showed lower in medium periods and higher in long periods of immersion. This indicates that the increase in pigment contents of leaves was ineffective in promoting P_n under long time immersion. Light saturation points under short time waterlogging (0–4 h) occurred at light intensities of 800–1000 μmol/m²/s, and at around 400 μmol/m²/s in long time treatments (8–12 h). Long periods of tidal immersion therefore significantly inhibited photosynthesis of mature leaves. Alcohol dehydrogenase and oxidase activity in roots both increased under longer immersion periods, suggesting that roots are sensitive to anaerobiosis under long term waterlogging. The activities of peroxidase and superoxide dismutase in mature leaves increased in 8 h and 10 h treatments, respectively. The content of malondialdehyde in mature leaves increased under long time treatments. Abscisic acid accumulation in mature leaves also had a sharp increase from 8 h to 12 h inundation. Even though the anti-oxidative enzymes were induced by waterlogging, this was not sufficient to protect the seedlings from senescence. The results suggested that K. candel seedlings completely tolerated tidal immersion by seawater up to about 8 h in each cycle, which matches the natural distribution of K. candel in inter-tidal zones of China.     

硫化氢对淹水平邑甜茶根系形态构型、叶片活性氧和光合特性的影响

以盆栽平邑甜茶为试材,分别将盆浸没于5种浓度硫化氢(H₂S)释放剂硫氢化钠(NaHS;0、0.02、0.05、0.1、0.2 mmol·L^-1)水溶液,以及添加有次牛磺酸(HT;H₂S清除剂)的NaHS水溶液中,10 d后调查根系形态构型、叶片活性氧生成和光合气体交换参数以及植株生长的变化.结果表明: 在淹水处理液中加入0.02～0.1 mmol·L^-1 NaHS,能增加土壤淹水时根系总长度、表面积、体积、直径、根尖数、分形维数、一级侧根数量和根系活力,降低叶片超氧阴离子(O₂^-·)生成速率和过氧化氢(H₂O₂)含量,提高叶片光合速率(P_n)、蒸腾速率(T_r)、气孔导度(g_s)、水分利用效率(WUE)和表观CO₂利用效率(CUE),还提高幼苗株高和植株生物量,其中NaHS在浓度0.1 mmol·L^-1 时作用效果最显著;当NaHS施用浓度达到0.2 mmol·L^-1或在0.1 mmol·L^-1 NaHS中加施次牛磺酸时,上述指标均降回到单纯淹水时的水平.这说明添加适量H₂S能够有效缓解土壤淹水对平邑甜茶根系生长的抑制,可以通过减少活性氧产生以及提高CUE,从而减轻淹水胁迫对叶片光合性能和植株生长的不利影响.     

Process of aerenchyma formation and reactive oxygen species induced by waterlogging in wheat seminal roots

The development and regulation of aerenchyma in waterlogged conditions were studied in the seminal roots of wheat. Evans blue staining and the first cell death position indicated that the cortical cell death began at the root mid-cortex cells in flooding conditions. Continuous waterlogging treatment caused the spread of cell death from the mid-cortex to the neighboring cells and well-developed aerenchyma was formed after 72 h. Meanwhile, the formation of radial oxygen loss barrier was observed in the exodermis owing to the induction of Casparian bands and lignin deposition. Analysis of aerenchyma along the wheat root revealed that aerenchyma formed at 10 mm from the root tip, significantly increased toward the center of the roots, and decreased toward the basal region of the root. In situ detection of radial oxygen species (ROS) showed that ROS accumulation started in the mid-cortex cells, where cell death began indicating that cell death was probably accompanied by ROS production. Further waterlogging treatments resulted in the accumulation of ROS in the cortical cells, which were the zone for aerenchyma development. Accumulation and distribution of H₂O₂ at the subcellular level were revealed by ultracytochemical localization, which further verified the involvement of ROS in the cortical cell death process (i.e., aerenchyma formation). Furthermore, gene expression analysis indicated that ROS production might be the result of up-regulation of genes encoding for ROS-producing enzymes and the down-regulation of genes encoding for ROS-detoxifying enzymes. These results suggest that aerenchyma development in wheat roots starts in the mid-cortex cells and its formation is regulated by ROS.     

涝渍对3个树种生长、组织孔隙度和渗漏氧的影响

为了了解落羽杉(Taxodium distichum)、乌桕(Sapium sebiferum)和美国山核桃(Carya illinoensis)等树种的耐涝机制, 采用盆栽模拟涝渍环境的试验方法, 设置了淹水、渍水和对照3个处理, 测定了一年生落羽杉、乌桕和美国山核桃实生苗的生长、组织孔隙度、根氧消耗等指标。结果表明, 涝渍处理抑制了落羽杉、乌桕和美国山核桃的生物量和生物量增量(渍水处理下落羽杉的生长得到了促进), 增加了3树种的根冠比, 从生物量和生物量增量下降幅度来评价, 落羽杉的耐涝性最强, 其次为美国山核桃。淹水和渍水处理下, 落羽杉、乌桕和美国山核桃的根、茎和叶中的组织孔隙度显著增加, 且随着处理时间的延长, 各器官的组织孔隙度有增加的趋势, 3个树种中, 落羽杉的根、茎和叶中的组织孔隙度均较其他2个树种高。淹水和渍水处理下, 移除茎明显增加了落羽杉、美国山核桃和乌桕的根的氧消耗, 表明涝渍处理增强了O₂在3个树种体内的运输并通过根系扩散到涝渍土壤中的能力, 并且随着处理时间的延长, 3个树种体内运输O₂并扩散到土壤中的能力有逐渐增强的趋势。因此, 涝渍环境总体上抑制了落羽杉、乌桕和美国山核桃等树种的生长, 但各树种为了适应这种生长环境, 形成了大量的通气组织, 从而导致各器官组织孔隙度的增加, 增强了O₂通过植物体运输到根系并扩散到土壤中的能力, 解决了根系及根际缺氧的矛盾。     

Identification of genes expressed in maize root cortical cells during lysigenous aerenchyma formation using laser microdissection and microarray analyses

? To adapt to waterlogging in soil, some gramineous plants, such as maize (Zea mays), form lysigenous aerenchyma in the root cortex. Ethylene, which is accumulated during waterlogging, promotes aerenchyma formation. However, the molecular mechanism of aerenchyma formation is not understood. ? The aim of this study was to identify aerenchyma formation-associated genes expressed in maize roots as a basis for understanding the molecular mechanism of aerenchyma formation. Maize plants were grown under waterlogged conditions, with or without pretreatment with an ethylene perception inhibitor 1-methylcyclopropene (1-MCP), or under aerobic conditions. Cortical cells were isolated by laser microdissection and their mRNA levels were examined with a microarray. ? The microarray analysis revealed 575 genes in the cortical cells, whose expression was either up-regulated or down-regulated under waterlogged conditions and whose induction or repression was suppressed by pretreatment with 1-MCP. ? The differentially expressed genes included genes related to the generation or scavenging of reactive oxygen species, Ca(2+) signaling, and cell wall loosening and degradation. The results of this study should lead to a better understanding of the mechanism of root lysigenous aerenchyma formation.     

Mechanisms of waterlogging tolerance in wheat – a review of root and shoot physiology

We review the detrimental effects of waterlogging on physiology, growth and yield of wheat. We highlight traits contributing to waterlogging tolerance and genetic diversity in wheat. Death of seminal roots and restriction of adventitious root length due to O₂ deficiency result in low root:shoot ratio. Genotypes differ in seminal root anoxia tolerance, but mechanisms remain to be established; ethanol production rates do not explain anoxia tolerance. Root tip survival is short‐term, and thereafter, seminal root re‐growth upon re‐aeration is limited. Genotypes differ in adventitious root numbers and in aerenchyma formation within these roots, resulting in varying waterlogging tolerances. Root extension is restricted by capacity for internal O₂ movement to the apex. Sub‐optimal O₂ restricts root N uptake and translocation to the shoots, with N deficiency causing reduced shoot growth and grain yield. Although photosynthesis declines, sugars typically accumulate in shoots of waterlogged plants. Mn or Fe toxicity might occur in shoots of wheat on strongly acidic soils, but probably not more widely. Future breeding for waterlogging tolerance should focus on root internal aeration and better N‐use efficiency; exploiting the genetic diversity in wheat for these and other traits should enable improvement of waterlogging tolerance.     

Dry matter production in relation to root plastic development, oxygen transport, and water uptake of rice under transient soil moisture stresses

Drought and waterlogging are important abiotic stresses negatively affecting plant growth and development. They are transiently recurring in rainfed lowlands and in water-saving system practicing intermittent irrigation. This study aimed to determine the contribution of plastic development and associated physiological responses of roots to shoot dry matter production under transient soil moisture stresses. To minimize effect of genetic confounding, a selected line (CSSL47) drawn from 54 chromosome segment substitution lines (CSSL) of Nipponbare (japonica type) carrying an overlapping chromosome segments of Kasalath (indica type), was used and compared with the recurrent parent Nipponbare. Under transient droughted-to-waterlogged (D–W) conditions, CSSL47 showed greater shoot dry matter production than Nipponbare. This was due largely to its greater root system development through high induction of aerenchyma formation. Consequently, aerenchyma development effectively facilitated the internal diffusion of oxygen (O₂) to the root tips under sudden waterlogged condition supporting rapid recovery of stomatal conductance, transpiration, and photosynthesis. Likewise, CSSL47 showed greater shoot dry matter production than Nipponbare under transient waterlogged-to-droughted (W–D) conditions. This was due to CSSL47’s greater root system development through more initiation of L type lateral roots that effectively maintained soil water uptake. This in turn sustained higher stomatal conductance, transpiration, and photosynthesis. Results implied that utilization of CSSLs could precisely reveal that root plastic development in response to transient soil moisture stresses contributed to the maintenance of shoot dry matter production.     

Root growth,aerenchyma development,and oxygen transport in rice genotypes subjected to drought and waterlogging

Soils under field conditions may experience fluctuating soil water regimes ranging from drought to waterlogging. The inability of roots to acclimate to such changes in soil water regimes may result in reduced growth and function thereby, dry matter production. This study compared the root and shoot growth, root aerenchyma development, and associated root oxygen transport of aerobic and irrigated lowland rice genotypes grown under well-watered (control), waterlogged, and droughted soil conditions for 30 days. The aerobic genotypes were as tolerant as the irrigated lowland genotypes under waterlogging because of their comparable abilities to enhance aerenchyma that effectively facilitated O₂ diffusion to the roots for maintaining root growth and dry matter production. Under drought, aerobic genotypes were more tolerant than the irrigated lowland genotypes due to their higher ability to maintain nodal root production, elongation, and branching, thus, less reduction in dry matter production. Aerenchyma was also formed in droughted roots regardless of genotypes, but was resistant to internal O₂ transport under O₂ deficiency. The ability of roots to resist temporal variations in drought and waterlogging stresses might have strong implications for the adaptation of rice growing in environments with fluctuating soil water regimes.