水分胁迫条件下草莓克隆分株间水分调控及其对光合功能的影响

以盆栽草莓(Fragaria×ananassa)为材料研究了水分胁迫下克隆植物草莓母株和子株间的水分调控机制及其与碳同化、光系统Ⅱ激发能分配的关系.实验材料分为匍匐茎连接和剪断两个大组,进行两步实验.第1步实验,对连接组和剪断组的所有母株控水,子株充分供水;4d后进入第2步实验,把连接组分为两小组,对其中一组充分供水子株开始控水,另一组保持不变.结果表明,土壤干旱引起母株叶片失水,并使其净光合速率和气孔导度显著降低.但是连接组中供水良好的子株能有效缓解缺水母株的水分胁迫.当供水良好的子株也开始受到干旱处理的时候,则会加剧与之相连母株的水分胁迫.受胁迫母株可以通过加强渗透调节能力和降低水势从相连子株获取水分.虽然土壤干旱会造成受胁迫母株叶片脱落酸(abscisic acid, ABA)含量的大幅度增加,但是与之相连子株的叶片ABA含量并没有增加;并且气孔导度与ABA变化趋势一致.(1)草莓母株和子株间的水分运输是由二者的水势差驱动的;(2)ABA不会通过匍匐茎在母株和子株间传递并影响相邻子株气孔导度;(3)在水分异质性较大情况下,生理整合可明显提高克隆系统的碳同化能力和光系统Ⅱ激发能利用效率.     

遮阴对疏叶骆驼刺叶形态和光合参数的影响

通过设置自然光与遮阴(60%自然光)两种光环境, 观测了遮阴60天后疏叶骆驼刺(Alhagi sparsifolia)叶形态、光合生理参数和脯氨酸(Pro)含量的变化。结果表明: 与自然光照下的叶片相比, 遮阴叶的比叶面积显著增大(p < 0.01), 比叶干重、比叶鲜重和叶片厚度明显降低(p < 0.01); 叶绿素(a + b)和类胡萝卜素含量有所增加, 其中叶绿素a含量增加显著(p < 0.01); 光补偿点、光饱和点和暗呼吸速率降低, 表观量子效率提高, 最大净光合速率明显增大, 光能利用效率高于自然光叶; 强光照下遮阴叶的净光合速率明显降低, 易发生光合光抑制现象。上述结果说明: 遮阴处理后, 疏叶骆驼刺在叶形态和光合生理上表现出对遮阴弱光条件的一定程度的驯化适应。另外, 遮阴叶片Pro的大量积累, 说明Pro在疏叶骆驼刺驯化适应弱光环境中起着重要作用。遮蔽环境下疏叶骆驼刺植株死亡率明显偏高, 表明塔克拉玛干沙漠南缘荒漠区的疏叶骆驼刺属于专性阳生植物不耐阴品种。     

疏叶骆驼刺根系对土壤异质性和种间竞争的响应

近年来, 植物根系对土壤异质性的响应和植物根系之间的相互作用一直是研究的热点。过去的研究主要是针对一年生短命植物进行的, 而且多是在人工控制的温室条件下进行的。而对于多年生植物根系对养分异质性和竞争的综合作用研究很少。该文对塔里木盆地南缘多年生植物疏叶骆驼刺(Alhagi sparsifolia)根系生长对养分异质性和竞争条件的响应途径与适应策略进行了研究, 结果表明: (1)在无竞争的条件下, 疏叶骆驼刺根系优先向空间大的地方生长, 即使另一侧有养分斑块存在, 其根系也向着空间大的一侧生长; (2)在有竞争的条件下, 疏叶骆驼刺根系生长依然是优先占领空间大的一侧, 但是竞争者的存在抑制了疏叶骆驼刺的生长, 导致其枝叶生物量和根系生物量都明显减少(p < 0.01), 而养分斑块的存在促进了疏叶骆驼刺根系的生长; (3)疏叶骆驼刺根系的生长不仅需要养分, 也需要足够的空间, 空间比养分更重要; (4)有竞争者存在的时候, 两株植物的根系都先长向靠近竞争者一侧的空间, 即先占据“共有空间”。研究结果对理解植物根系觅食行为和植物对环境的适应策略有重要意义。     

间隔子长度对美丽箬竹克隆分株水分生理整合效应的影响

克隆植物分株间隔子长度(即相邻分株间距离)可能对生理整合产生显著的影响,其抗氧化系统与光合色素的变化可能影响植物对水分异质性的响应能力。该研究以克隆植物美丽箬竹(Indocalamus decorus)为材料,设高[H,(90±5)%]、中[M,(60±5)%]和低[L,(30±5)%]3个基质相对含水率和2个水势梯度(H-L和M-L),测定了3种间隔子长度(10、25和40cm)根状茎相连克隆分株叶片抗氧化酶活性、相对电导率以及丙二醛(MDA)、可溶性蛋白质和光合色素含量,分析美丽箬竹水分生理整合的水分梯度效应、间隔子长度效应与时间效应。结果显示:(1)随处理时间的延长,分株叶片CAT活性、相对电导率及MDA含量均先升高后降低,而叶片SOD活性则与之相反;叶片POD活性和光合色素含量总体上呈"N"型变化,而可溶性蛋白质含量的变化趋势与POD活性呈倒"N"型。(2)水分梯度和间隔子长度均会对美丽箬竹分株间水分生理整合产生影响,随分株间隔子长度增大和水势梯度差减小,低水势受体分株叶片抗氧化酶活性、可溶性蛋白质含量和光合色素含量明显降低,而其相对电导率和丙二醛(MDA)含量明显升高;供体分株水势越高,分株间间隔子越长,供体分株叶片抗氧化酶活性、可溶性蛋白质和光合色素含量越高,而其相对电导率和MDA含量明显降低。研究表明,在异质水分环境下美丽箬竹克隆系统发生了从高水势向低水势分株的水分转移,分株间水势梯度差越大、间隔子长度越短,水分传输的整合强度越高,低水势受体分株获益越明显;随着处理时间的延长,分株间水分生理整合强度在处理前期增强,而在处理后期减弱,反映出供体分株与受体分株间损耗-受益在时间序列上是有变化的,且处理前期损耗-受益表现更为明显。     

极端干旱区疏叶骆驼刺叶绿素荧光对人工水分干扰的响应特征

选取塔里木河下游天然植被恢复示范区的疏叶骆驼刺(Alhagi sparsifolia Shap.)为研究对象,测定每年引水灌溉2次(每次0.42m3/m2)、1次和不灌溉(CK)处理下疏叶骆驼刺的实际光化学量子产量(ΦPSⅡ)、电子传输速率(ETR)和光化学猝灭系数(qP)等叶绿素荧光参数及其叶水势变化,探讨疏叶骆驼刺对人工水分干扰的叶绿素荧光响应特征。结果表明:(1)随着灌溉量的减少,疏叶骆驼刺叶水势呈显著降低的趋势,并在CK下达到最低。(2)同期疏叶骆驼刺qP、ΦPSⅡ、ETR、最大光量子产量(Fv/Fm)、叶绿素含量和光饱和点均随着灌溉量的减少呈先增加后降低的趋势,非光化学淬灭系数(NPQ)和调节性能量耗散(YNPQ)则呈先降低后增加的趋势。(3)与每年1次灌溉量处理相比,不灌溉和每年2次的灌溉量处理下疏叶骆驼刺发生了光抑制,光能捕获效率与光化学反应能量下降,热耗散能力提高。研究认为,灌溉量过高(每年2次,0.84m3/m2)或不灌溉均会限制疏叶骆驼刺光化学效率和光和活性,适时适量的(春季灌水1次,0.42m3/m2)水分补给更有利于疏叶骆驼刺适应干旱胁迫并维持正常光合生长。     

高寒矮生嵩草草甸主要植物物候特征对养分和水分添加的响应

植物物候特征对环境条件的季节和年际变化具有较强的指示作用, 因此研究植物物候特征对环境条件变化的响应, 对理解植物和环境之间的相互作用关系、植物的适应机制和生存策略, 以及应对全球变化都具有重要的意义。该研究基于2009-2011年高寒矮生嵩草(Kobresia humilis)草甸养分水分控制实验的植物物候观测数据资料, 采用巢式方差分析、物候指数和聚类分析方法, 开展了高寒矮生嵩草草甸主要植物物候特征对养分和水分添加的响应研究。结果表明: (1)养分添加处理之间植物返青期和枯黄期均无显著差异, 但养分添加中氮磷处理对主要物种作用较明显, 使莎草科、禾本科、杂类草主要代表植物的返青期和枯黄期推迟。(2)增雪处理效应明显, 主要优势物种无论是何种养分添加, 在增雪处理后均表现出花期物候提前的趋势(p < 0.01), 同时增雪处理使杂类草植物返青期显著提前(p < 0.05)。增水处理对植物的作用效果并不一致, 其中垂穗披碱草(Elymus nutans)和双柱头藨草(Scirpus distigmaticus)的枯黄期显著推迟(p < 0.05), 而杂类草枯黄期提前。(3)养分添加后, 不同物种的物候特征表现出显著差异(p < 0.01), 例如雪白委陵菜(Potentilla nivea)枯黄期显著推迟(p < 0.05), 而双柱头藨草的枯黄期显著提前(p < 0.05), 但物种对养分添加响应的差异以植物类群为单位, 禾本科植物表现为返青期推迟, 而莎草科植物表现为返青期提前。(4)矮生嵩草草甸主要植物营养生长期与果后营养期持续天数之间呈负相关关系, 主要植物物候特征经聚类分析可以分为3个类群, 3个类群经氮磷钾、钾和氮钾三个养分添加处理后植物物候特征变化较大。研究表明, 高寒矮生嵩草草甸植物物候特征在物种水平响应和水分添加后的响应表现出较大差异, 而对养分添加的响应不显著。     

Effects of nitrogen addition on clonal integration between mother and daughter ramets of Moso bamboo: a 13C-CO2 pulse labeling study

用¹³C-CO₂脉冲标记法研究氮添加对毛竹母子分株克隆整合的影响 相连分株间的资源共享(即克隆整合)是克隆植物的显著特征。克隆整合使毛竹(Phyllostachys pubescens)对多种环境条件具有较强的适应性。但毛竹通过克隆整合获得性能提升的机制尚不明确。本 研究区分并分析了毛竹光合碳的顶向整合和基向整合,以探究毛竹克隆片段如何在土壤氮异质性条件下 提高整体收益。本研究以由两株不同年龄毛竹分株组成 的克隆片段为研究对象,分株间通过根状茎连接。 每个分株设置氮添加或无氮添加两种处理,并通过单株母株或子株的¹³C-CO₂脉冲标记对光合碳的顶向整合和基向整合进行区分。研究结果表明,子株氮添加显著促进了光合碳由母株向子株的顶向整合,无论母株是否有氮添加,均向氮添加子株转运更多的光合碳。转运自无氮添加母株的光合碳主要分配至氮添加子株的 叶片,而转运自氮添加母株的光合碳则主要分配至根系。与无氮添加处理的子株相比,氮 添 加子株转运更多的光合碳至母株,且该过程不受母株氮添加的影响。转运自氮添加子株的光合碳主要 分配至无氮添加母株的根系和氮添加母株的叶片。这些结果说明,母株优先将更多的资源投入到具有高 养分可利用性的子株,随后子株作为更高效的资源采集点,根据母株的养分状况针对富集资源进行获取。通过这种方式,克隆植物可以降低资源获取成本并提高资源获取效率,最大限度地提升其整体表现。     

干旱及复水条件下接种AMF和根瘤菌对疏叶骆驼刺根系生长的影响北大核心CSCD

疏叶骆驼刺为塔里木河下游优势草本植物,对下游地区防风固沙,涵养水源具有重要的生态价值。该试验以疏叶骆驼刺为研究对象,设定正常水分(土壤相对含水量70%±5%)、干旱胁迫(土壤相对含水量20%±5%)和复水处理(干旱胁迫60 d后恢复至正常水分)3个水分梯度,以及单接种丛枝菌根真菌、单接种根瘤菌、双接种丛枝菌根真菌+根瘤菌和不接种4组接种处理,分析不同水分条件下各接种处理对疏叶骆驼刺根系生长的影响。结果表明:(1)双接种丛枝菌根真菌+根瘤菌处理的疏叶骆驼刺根系AMF侵染率在干旱胁迫、复水条件下均显著降低,且低于单接种AMF处理。(2)随着正常水分→干旱胁迫→复水的水分变化,双接种处理疏叶骆驼刺根系根瘤数量先降低后增加,复水后显著高于单接种根瘤菌处理。(3)双接种处理扩大了疏叶骆驼刺的根系吸收范围,提高了根系的吸收能力,并随着正常水分→干旱胁迫→复水的水分变化,呈现先降低后增加的变化趋势。(4)双接种处理显著提高了疏叶骆驼刺根系SOD和POD活性,并随着正常水分→干旱胁迫→复水的水分变化而逐渐升高。研究发现,双接种AMF+根瘤菌处理可以显著促进疏叶骆驼刺根系的生长,增强其抗逆性,而干旱胁迫会降低AMF和根瘤菌的协同促进作用,复水后双接种AMF+根瘤菌处理的疏叶骆驼刺能及早地做出响应,对其根系生长表现出一定的补偿效应。     

美丽箬竹水分生理整合的分株比例效应——基于叶片抗氧化系统与光合色素

生理整合是克隆植物实现资源共享, 增强对异质生境适应能力的重要手段。其中, 水分生理整合是克隆植物最为重要的生理整合, 解析竹子水分生理整合特征对于竹林水分科学管理具有重要意义。该研究以分株地下茎相连的美丽箬竹(Indocalamus decorus)盆栽苗为试验材料, 设置2个盆栽基质相对含水率(高水势(90% ± 5%)和低水势(30% ± 5%))和5个分株比例(1:3、1:2、1:1、2:1、3:1, 高水势分株与低水势分株数量比值, 地下茎相连的分株总数12株)处理。处理后15、30、45、60天分别取不同处理的克隆分株成熟叶测定抗氧化酶活性、相对电导率和丙二醛含量、可溶性蛋白质含量、光合色素含量, 分析基于分株比例的美丽箬竹水分生理整合方向、强度和效率的变化规律。结果表明: 在异质水分条件下, 美丽箬竹分株间存在着从高水势供体分株向低水势受体分株进行水分转移的生理整合作用, 并随着分株比例的增大, 整合强度增强, 受体分株获益提高, 供体分株耗损增大。随着处理时间的延长, 处理前期分株间水分生理整合强度增强, 处理后期整合强度减弱, 反映出供体分株与受体分株间耗-益在时间序列上是有变化的, 处理前期耗-益更为明显。研究表明克隆系统分株比例对竹子水分生理整合有重要影响, 分株间水分梯度差是水分传导的潜在驱动力, 决定水分生理整合方向、强度和效率的是分株间水分供需关系。     

水分异质性影响两种根茎型克隆植物赖草和假苇拂子茅的水分存储能力

水分在自然系统中呈异质性分布。有关水分异质性对克隆植物生长、形态和生理影响的研究已有大量的工作, 但是水分异质性对克隆植物存储能力, 尤其是水分存储能力影响的研究却十分缺乏。该文将两种根茎型克隆植物赖草(Leymus secalinus)和假苇拂子茅(Calamagrostis pseudophragmites)进行水分异质性和同质性实验处理, 探讨水分异质性对克隆植物水分存储能力、生长和形态的影响。在异质性水分处理下, 两种克隆植物的间隔子、枝和根的含水量均显著增加。两种克隆植物对水分异质性分布的适应策略有所不同, 赖草通过降低单个克隆分株的生长、提高芽的数量以应对水分异质性, 而假苇拂子茅通过增强整个分株种群的地下部分(根状茎、根和芽)生长来应对水分资源的异质性分布。水分储存能力的增强可以提高克隆植物适应水分异质性的能力。     

Clonal integration enhances flood tolerance of Spartina alterniflora daughter ramets

Recently, considerable attention has been paid to the invasion of the clonal plant Spartina alterniflora into coastal wetlands at lower elevations. In this experiment, we tested whether clonal integration improved flood tolerance in S. alterniflora daughter ramets. Daughter ramets at two growth stages (young and old ramets) were flooded to water levels of 0, 9 and 18 cm above the soil surface, and the rhizomes between mother and daughter ramets were either severed or left intact. Biomasses of connected ramets grown in controls or in shallow and deep water treatments were 119%, 108% and 149% higher in the old ramet group than those of severed ramets, respectively, whereas they were 3.0, 3.3 and 11.2 times higher in the young ramet group, respectively. At the end of the experimental period, the shoot height, connected with young ramets, in shallow and deep water treatments increased by 19% and 26%, respectively, over that in the control treatments, whereas the old ramets increased by 11% and 39%, respectively. In contrast, the shoot height of the severed young ramets was 27% and 26% lower in shallow and deep water treatments than in the control treatment, respectively. However, the shoot height of the severed old ramets remained constant with increasing water depth. We conclude that clonal integration enhances the flood tolerance of S. alterniflora daughter ramets, and the trait of clonal integration plays more important roles in severe flooding stress conditions and at early growth stages.     

Clonal integration in Ranunculus reptans: by‐product or adaptation?

We studied fitness consequences of clonal integration in 27 genotypes of the stoloniferous herb Ranunculus reptans in a spatially heterogeneous light environment. We grew 216 pairs of connected ramets (eight per genotype) with mother ramets in light and daughter ramets in shade. In half of the pairs we severed the stolon connection between the two ramets at the beginning of the experiment. During the experiment, 52.7% of the ramet pairs with originally intact connection physically disintegrated. We detected significant variation among genotypes in this regard. Survival of planted ramets was 13.3% higher for originally connected pairs. Moreover, there was significant variation among genotypes in survival, in the difference in survival between plant parts developing from mother and daughter ramets, and in the effect of integration on this difference. In surviving plants connection between ramets decreased size differences between mother and daughter parts. Variation among genotypes was significant in growth and reproduction and marginally significant in the effect of physiological integration on growth and reproduction. Connected daughter ramets had longer leaves and internodes than daughters in severed pairs indicating that integration stimulated plant foraging in both the vertical and the horizontal plane. Observed effects of integration on fitness components in combination with genetic variation in maintenance and effects of connection indicate that clonal integration in R. reptans has the capability to evolve, and therefore suggest that clonal integration is adaptive. If genetic variation in integration is common, future studies on clonal integration should always use defined genetic material and many clones to allow extrapolation of results to population and wider levels.     

Does clonal integration improve competitive ability? A test using aspen (Populus tremuloides [Salicaceae]) invasion into prairie

Many clonal plants consist of many connected individual ramets, allowing them to share water and nutrients via physiological integration. Integration among ramets may also improve the ability of clonal plants to tolerate abiotic stress or improve the competitive ability of individual ramets. Here I use a field experiment to determine whether clonal integration improves ramet performance for a widespread clonal tree species invading into native prairie. Aspen (Populus tremuloides) dominates the southern treeline in western Canada, has long-lived belowground connections between mother and daughter ramets, and reproduces vegetatively via resprouting rhizomes after disturbance. I applied two competition treatments (neighbors present or absent) and two clonal integration treatments (belowground rhizomes between mother and daughter ramets either severed or left intact) to 12 replicate Populus daughter ramets at each of three sites. Neighbors improved the survivorship of Populus ramets by 25-35% after 2 yr, but decreased growth by ～20%. Clonal integration tended to improve ramet survival and growth, but these trends were often not significant. Clonal integration did not alter the effects of competition from neighboring vegetation, suggesting that connections between ramets do not necessarily improve the competitive ability of Populus invading into native prairie.     

Clonal integration affects growth and sediment properties of the first ramet generation,but not later ramet generations under severe light stress

克隆整合影响严重光胁迫下第一分株世代的生长和沉积物特征但不影响 后续分株世代的生长和沉积物特征 克隆整合通过缓冲环境压力和提高资源获取效率使克隆植物受益。然而,在一个克隆系统中,受益于克隆整合的连接分株世代的数量很少受到关注。我们进行了一个盆栽实验来评估沉水植物苦草 (Vallisneria natans)克隆系统内的生理整合程度,该克隆系统由一个母株和3个依次连接的后代分株组成。 母株生长在正常光照下,而后代分株被严重遮荫。母株与后代分株间的匍匐茎被切断或保持连接,但3个后代分株之间的连接仍然存在。与遮荫的后代分株连接时,苦草未遮荫的母株的光合能力显著增强,但其生物量积累大大减少。克隆整合显著增加了第一分株世代(相邻分株)的生物量积累和土壤的碳氮可用性、胞外酶活性和微生物生物量,但没有增加后续分株世代的这些特征。我们的结果表明,在严重光胁迫下,来自苦草母株的支持可能仅限于克隆系统中相邻的后代分株,这暗示着一个分株世代的效应。我们的结果有助于更好地理解克隆植物的层次结构和分段化。这些发现表明克隆整合程度在分株种群的生态相互作用中起着至关重要的作用。     

The effects of clonal integration on morphological plasticity and placement of daughter ramets in black locust (Robinia pseudoacacia)

We studied the field response of Robinia pseudoacacia L. to light, total soil nitrogen, available soil phosphorus and soil pH. Results indicated that there was very strong clonal integration between mother and daughter ramets. Mother ramets can provide nitrogen and phosphorus to daughter ramets sufficient for their continued growth through strong clonal integration, but cannot provide enough photosynthate. With clonal integration, soil nitrogen and phosphorus availability had no effect on biomass allocation to roots, number of ramets and length of connection roots. Biomass allocation to roots increased markedly and responded to nitrogen and phosphorus availability, when the connections were severed. Light had a significant effect on the percent of biomass allocation to leaves and number of ramets, but no effect on the length of connection roots. Daughter ramets allocated more resources to leaves, and clones placed more daughter ramets in high light patches than in low light patches. Soil pH had a significant effect on ramet number and connection root length. Clones concentrated in alkaline patches and escaped from acid patches through selective placement of daughter ramets and changing the length of connection roots. We suggest that the clonal integration may be very strong and provide sufficient soil resources to daughter ramets, then affect the daughter ramets’ morphology and placement, if the size of a specific ramet is significantly larger than the other ramets in an arbor clone.     

Effects of the loss of clonal integration on four sedges that differ in ramet aggregation

Although clonal growth is a dominant mode of plant growth in wetlands, the importance of clonal integration, resource sharing among ramets, to individual ramet generations (mother and daughter) and entire clones of coexisting species has not been well investigated. This study evaluated the significance of clonal integration in four sedge species of varying ramet aggregations, from clump-forming species (Clumpers –Carex sterilis, Eleocharis rostellata), with tightly aggregated ramets (rhizomes<1cm), to runner species (Runners –Schoenoplectus acutus, Cladium mariscoides), with loosely aggregated ramets. We manipulated clonal integration by either severing connections between target mother and daughter ramets or leaving connections intact, and then planted them in an intact neighborhood of a fen in Michigan, USA. We measured growth parameters of original and newly produced ramets over two growing seasons and conducted a final biomass harvest, to address four hypotheses. First, we expected integrated clones to accumulate more biomass than severed clones. However, final clone-level biomass and ramet production were the same for both treatments in all species although severing initially stimulated ramet production by Schoenoplectus and produced a more compact ramet aggregation in Cladium. Second, we hypothesized that mother ramets would experience a cost of integration, through reduced ramet or biomass production, while daughters would experience a benefit, through increased resource availability from mothers. Mother ramets of Cladium suffered a cost from integration, while Schoenoplectus mothers suffered a slight cost and Carex daughters saw a slight benefit. Finally, we hypothesized that integration would be more active in runner species than in clumper species. Indeed, we documented more active integration in runners than clumpers, but none of the study species were dependent upon integration for growth or survival once daughter ramets were established with their own roots and shoots. This study demonstrates that integration between established ramets may not be the most important advantage to clonal growth in this wetland field site. The loss of integration elicited varied responses among coexisting species in their natural habitat, somewhat but not completely related to their growth form, suggesting that a combination of plant life history traits contributes to the dependence upon clonal integration among established ramets of clonal species.     

Water translocation between ramets of strawberry during soil drying and its effects on photosynthetic performance

To explore the mechanisms underlying water regulation in clonal plants and its effects on carbon assimilation under water stress, we studied the responses of water status, gas exchange and abscisic acid (ABA) contents to water stress in leaves of pairs of strawberry ramets that consist of mother and daughter ramets. There was a greater decrease in photosynthetic rates (P_n) and stomatal conductance (G_s) in the disconnected mother ramets than the connected mother ramets upon exposure to water stress, indicating that water stress in mother ramets was alleviated by water translocation from the well‐watered daughter ramets. Conversely, the connected mother ramets displayed enhanced symptoms of water stress when the connected daughter ramets were exposed to water deficit. The mother ramets had lower water potential (ψ_w) due to their stronger osmotic adjustment than in well‐watered daughter ramets; this resulted in water flow from the connected daughter ramets to mother ramets, thus alleviating water stress of mother ramets. During soil drying, there was a striking increase in ABA concentrations in leaves of the disconnected mother ramets, whereas leaf bulk ABA was much lower in the connected and water‐stressed mother ramets than that in the drought‐affected mother ramets in the disconnected group. In this study, though G_s was linearly correlated with leaf bulk ABA and ψ_w, G_s in water‐stressed mother ramets in disconnected group exhibited less sensitivity to the variation in leaf bulk ABA and ψ_w than that in connected and water‐stressed mother ramets. Taken together, these results indicate that: (1) the flux of water translocation between the connected ramets is determined by a water potential gradient; (2) water translocation between connected ramets helps to keep sensitivity of G_s to ABA and ψ_w in drought‐affected ramets, thereby benefit to effectively maintain the homeostasis of leaf water status and (3) the improvements in P_n in water‐stressed ramets due to water translocation from well‐watered ramets suggest the advantages of physiological integration in clonal plants in environments with heterogeneous water distribution.     

Effects of inoculation with native arbuscular mycorrhizal fungi on clonal growth of Potentilla reptans and Fragaria moschata (Rosaceae)

Many clonal plants live in symbiosis with ubiquitous arbuscular mycorrhizal (AM) fungi, however, little is known about their interaction with respect to clonal reproduction and resource acquisition. The effects of arbuscular mycorrhiza on the growth and intraclonal integration between ramets of two stoloniferous species were studied experimentally in a nutritionally homogenous soil environment. Two species coexisting at the same field site, Potentilla reptans and Fragaria moschata, were selected as model plants for the study. Pairs of their ramets were grown in neighbouring pots with each ramet rooted separately. Four inoculation treatments were established: (1) both mother and daughter ramets remained non-inoculated, (2) both ramets were inoculated with a mixture of three native AM fungi from the site of plant origin, (3) only mother or (4) daughter ramet was inoculated. The stolons connecting the ramets were either left intact or were disrupted. Despite the consistent increase in phosphorus concentrations in inoculated plants, a negative growth response of both plant species to inoculation with AM fungi was observed and inoculated ramets produced fewer stolons and fewer offspring ramets and had lower total shoot dry weights as compared to non-inoculated ones. A difference in the extent of the negative mycorrhizal growth response was recorded between mother and daughter ramets of P. reptans, with daughter ramets being more susceptible. Due to AM effect on ramet performance, and thereby on the source-sink relationship, inoculation also significantly influenced biomass allocation within clonal fragments. Physiological integration between mother and daughter ramets was observed when their root systems were heterogeneous in terms of AM colonization. These results hence indicate the potential of mycorrhizal fungi to impact clonal growth traits of stoloniferous plant species, with possible consequences for their population dynamics.     

Physiological Integration Ameliorates Negative Effects of Drought Stress in the Clonal Herb Fragaria orientalis

Clonal growth allows plants to spread horizontally and to establish ramets in sites of contrasting resource status. If ramets remain physiologically integrated, clones in heterogeneous environments can act as cooperative systems - effects of stress on one ramet can be ameliorated by another connected ramet inhabiting benign conditions. But little is known about the effects of patch contrast on physiological integration of clonal plants and no study has addressed its effects on physiological traits like osmolytes, reactive oxygen intermediates and antioxidant enzymes. We examined the effect of physiological integration on survival, growth and stress indicators such as osmolytes, reactive oxygen intermediates (ROIs) and antioxidant enzymes in a clonal plant, Fragaria orientalis, growing in homogenous and heterogeneous environments differing in patch contrast of water availability (1 homogeneous (no contrast) group; 2 low contrast group; 3 high contrast group). Drought stress markedly reduced the survival and growth of the severed ramets of F. orientalis, especially in high contrast treatments. Support from a ramet growing in benign patch considerably reduced drought stress and enhanced growth of ramets in dry patches. The larger the contrast between water availability, the larger the amount of support the depending ramet received from the supporting one. This support strongly affected the growth of the supporting ramet, but not to an extent to cause increase in stress indicators. We also found indication of costs related to maintenance of physiological connection between ramets. Thus, the net benefit of physiological integration depends on the environment and integration between ramets of F. orientalis could be advantageous only in heterogeneous conditions with a high contrast.