雌雄异株植物对环境胁迫响应的性别差异研究进展

从温度、CO2、水分以及生境变化4个方面综述了雌雄异株植物的雌雄个体在环境胁迫下出现的不同生理、生态和生化差异.温度胁迫将导致雌雄个体在气孔导度、净同化作用、耐冻性形成以及ABA含量等方面出现不同的生理响应,响应大小受限于外界条件.雄株植物的净光合速率高于雌株,而且随着CO2浓度的增高,二者的光合作用受到明显的促进,生物量均显著增加.水分胁迫使雌雄个体在干物质积累、净光合速率、蒸腾速率、水分利用效率和碳同位素值等生理指标以及部分形态指标呈现显著差异.多数植物的雄株个体在干旱环境中具有较高的水分利用效率,对水分胁迫的抗性更大.不同生境中雌雄异株植物雌雄个体的适应性各不相同.在土壤干燥、养分贫瘠、海拔较高、坡度较大的生境中,雄性植株生长良好,数量较多;而在湿润、肥沃、低海拔或低洼的生境中,雌性植株生长良好.     

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。     

克隆植物沙拐枣的母株和分株对风蚀沙埋的生理生态响应

以中国荒漠区优良的防风固沙克隆灌木沙拐枣为对象,研究了长期风蚀、沙埋环境下沙拐枣母株和克隆分株的同化枝对环境异质性的响应。结果发现:(1)风蚀母株、风蚀分株的叶绿素含量、净光合速率、气孔导度、蒸腾速率、胞间CO2浓度和水分利用效率只有沙埋分株的一半左右,导致同化枝的长度、数量、簇数也仅是沙埋分株的一半,而且风蚀母株的果实宽和果实长也都最小,但浅沙埋有利于沙拐枣的生长和繁殖,表明严峻的风蚀对母株和分株的生长与繁殖都产生了胁迫,但浅沙埋有利于沙拐枣的生长和繁殖。(2)风蚀母株倒伏后同化枝的形态特征是基部优于中部优于顶部,表明严峻风蚀下母株的死亡是从顶部-中部-底部逐渐舍弃的过程。(3)母株的全部根系以及风蚀水平根全部裸露在外但依然能够存活,间接证明沙拐枣克隆整合的方向性——不仅可在分株间进行传递,分株-母株间也可进行传递,否则遭受严峻风蚀胁迫的母株和克隆分株会直接死亡。本研究结果为沙拐枣克隆生长对风沙环境的生态适应机制提供了基础,也是对植物克隆生态学在自然异质环境中研究缺乏的有效补充。     

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。     

养分异质条件下结缕草克隆分株生长及光合作用的生理整合

通过对连接和断开的分株进行不同养分处理,研究养分异质条件下结缕草克隆分株生长及光合作用的变化.结果表明:在养分异质条件下,处于中、高养分水平的母株可以提高与其相连子株的叶长、叶宽、根质量、叶质量、光合速率、气孔导度、蒸腾速率和水分利用效率,高养分条件下分别提高16.0%、8.3%、24.4%、58.1%、30.3%、54.0%、9.2%和21.9%,降低根冠比和胞间CO2浓度,在高养分下分别降低21.6%和31.5%;处于中、高养分水平的子株对与其相连母株的生长及光合特性没有显著影响.在养分异质条件下,结缕草母株对子株存在生理整合,养分梯度越大,整合强度越强.结缕草克隆子株可以从母株获益,但母株不能从子株获益,子株是生理整合单向获益者.     

孑遗植物银杏(Ginkgo biloba L.)伴性光合生理特征与进化生态

银杏类是一类古老的雌雄异株植物,目前仅存单科单属.由于银杏在系统发育等方面的独特地位,吸引了科学家从不同方面进行了广泛的研究.在适宜的条件下,测定了生长在野外环境下的成年银杏的雌雄个体的光合特性.研究结果表明,银杏的雌雄植株对光具有相同的表现趋势,光饱和点,光补偿点,光下呼吸速率等均没有明显差异.但是雌性银杏的净光合速率明显大于雄性银杏的净光合速率.这种差异可能与雌性个体在繁衍后代时需要投入更多的能量有关.研究也表明,在与其它的裸子植物和被子植物比较时,银杏的光合能力并没有明显的弱势,因此光合能力可能并不是银杏在第三纪分布受限的直接原因.     

孑遗植物银杏(Ginkgo biloba L.)伴性光合生理特征与进化生态

银杏类是一类古老的雌雄异株植物,目前仅存单科单属。由于银杏在系统发育等方面的独特地位,吸引了科学家从不同方面进行了广泛的研究。在适宜的条件下,测定了生长在野外环境下的成年银杏的雌雄个体的光合特性。 研究结果表明,银杏的雌雄植株对光具有相同的表现趋势,光饱和点,光补偿点,光下呼吸速率等均没有明显差异。但是雌性银杏的净光合速率明显大于雄性银杏的净光合速率。这种差异可能与雌性个体在繁衍后代时需要投入更多的能量有关。研究也表明,在与其它的裸子植物和被子植物比较时,银杏的光合能力并没有明显的弱势,因此光合能力可能并不是银杏在第三纪分布受限的直接原因.     

分株比例对异质光环境下美丽箬竹克隆系统光合生理的影响

分株数量或生物量比例差异会明显影响克隆系统对资源异质性环境的生态适应性, 地下茎木质化、连接稳固的竹类植物在生长过程中相连克隆分株通常会生活在异质光环境中, 但其叶片光合生理特性对异质光环境的响应及其分株比例效应则未见报道。该研究以地下茎相连的美丽箬竹(Indocalamus decorus)克隆系统为实验材料, 设置2个遮光率(分别为50% ± 5%和75% ± 5%)和3个分株比例(遮光与未遮光分株比例分别为1:3、2:2、3:1)处理, 分株数量为4株。分别测定了遮光处理后30、90、150天遮光和未遮光分株叶片光响应特征、气体交换参数、光合色素含量, 分析了异质光环境下美丽箬竹光合生理的变化规律。结果显示: 分株比例对美丽箬竹光合生理有显著影响, 且其与遮光、处理时间交互作用显著。美丽箬竹克隆系统遮光分株比例越大, 即遮光相对分株数量越多, 其表观量子效率(AQE)、光饱和点、最大净光合速率(P_{n max})、净光合速率(P_n)、气孔导度(G_s)、蒸腾速率(T_r)、水分利用效率越大, 光补偿点、暗呼吸速率越小, 其光合效率越高, 光能利用能力越强, 而与之相连的未遮光分株则相反; 随遮光分株比例的增大, 遮光分株叶片叶绿素a、叶绿素b含量呈先升高而后下降的变化趋势, 类胡萝卜素(Car)含量则持续下降, 而与之相连的未遮光分株叶片光合色素含量则呈下降趋势; 遮光率提高, 相同分株比例美丽箬竹克隆系统遮光分株叶片AQE、P_{n max}、P_n、G_s、胞间CO₂浓度(C_i)和光合色素含量总体升高, 而与之相连的未遮光分株叶片P_{n max}、C_i以及Car含量则总体下降。研究结果表明异质光环境下, 遮光分株光合效率和弱光利用能力明显增强, 而未遮光分株则相反, 克隆系统内分株间发生了明显的克隆分工, 且2:2、3:1分株比例克隆系统较1:3分株比例对异质光环境具有更好的适应能力。美丽箬竹克隆系统可通过差异性调节分株光合生理特性和光合色素含量, 提高遮光分株光能利用和光合效率来适应异质光环境, 以提高克隆系统的适合度。     

空心莲子草性可塑性影响下的克隆整合

植物的生活史由其有性生殖构件和营养体构件相互作用共同完成,克隆整合作为克隆植物的重要特征,其与有性生殖特征的相互作用关系却所知很少。该研究通过同质园种植实验,分析了空心莲子草的分株表型、生理、性别等与克隆整合的关系。结果表明:(1)克隆整合以及分株间是否连接对空心莲子草的表型特征、气体交换等生理性状和性别特征均有显著影响。(2)克隆整合显著缩小了雌雄同花和雄蕊心皮化两种性别植株间表型特征的差距,后代的性别特征与营养体表型特征显著相关。(3)在贫瘠的沙土基质中克隆整合明显增加了空心莲子草的营养体生长特征和气体交换等光合生理指标,但这种增加在富含有机质的塘泥基质中不明显。(4)居于不同土壤基质分株间的联系会减少分株表型特征和气体交换对生长环境的响应,并保持母体性别特征不受环境的影响,但单独居于沙土或塘泥单一土壤基质的分株性别特征却因受到环境影响而改变。因此,克隆整合有利于空心莲子草性别特征的稳定。     

百里香无性系的克隆生长特征

植物克隆生长及其与生态适应性的关系是当今植物种群生态学研究的热点和前沿课题,但目前小半灌木克隆生长的研究开展不多。百里香(Thymus serpyllum var. asiaticus)是一种具有地面匍匐茎的草本状小半灌木,可在土壤侵蚀剧烈、基岩大面积裸露的砒砂岩区形成百里香单优群落,在维持生态系统稳定方面具有重要的生态学作用。皇甫川流域是砒砂岩大面积分布的典型区域,在这一地区对百里香无性系的克隆生长进行研究,不仅具有重要的学术价值,而且在生态环境建设方面也具有一定的现实意义。在皇甫川流域选择含三级分株的百里香无性系,对其各级分株的总生物量、各构件生物量及数量、各构件生物量占总生物量的百分比及其月变化进行了研究。结果表明: 1)母株与子代相比,在总生物量、构件生物量及数量上占有绝对优势,而且具有体型大、结构复杂的特点; 2)对生物量分配格局的研究显示,母株根的生物量在总生物量中所占的比例最大,其叶所占的比例较低。子代叶的生物量在总生物量中所占的比例最大,其根所占的比例较低;3)不同级别分株在生物量分配上的差异,揭示了相互连接的分株在功能上的差别,母株可能更侧重于养分和水分的吸收,子株则更侧重于光合生产;4)构件枝、茎、花生物量分配比月变化显示,子1代各构件的生长规律与母株的基本一致,子2代与母株和子1代的相比差异较大,分析认为这可能是分株间不同程度的生理整合作用造成的结果。     

资源异质性环境中三种匍匐茎草本植物克隆内资源共享和分株功能特化

研究了3种来自中国北方林下、草地和碱化草甸匍匐茎型克隆草本植物绢毛匍匐委陵菜 (Potentilla reptans L. var. sericophylla Franch.)、鹅绒委陵菜 (P. anserina L.) 和金戴戴 (Halerpestes ruthenica (Jacq.) Qvcz.) 对由高光照低养分斑块和低光照高养分斑块组成的资源交互斑块性生境的适应性对策.当生长于高光照低养分条件下分株 (HL分株) 与生长于低光照高养分条件下分株 (LH分株) 之间的匍匐茎连接时, 3种克隆植物HL分株、LH分株以及整个分株对系统 (HL分株 + LH分株) 的生物量均得到显著提高.同时, LH分株根冠比显著增加, 而HL分株根冠比显著下降.这表明, 当互连分株置于由低光照高养分斑块和高光照低养分斑块组成的异质性环境中时, 3种植物克隆分株均发生了环境诱导的功能特化.克隆内资源共享以及克隆内不同分株的功能特化有利于整个分株系统对局部丰富资源的获取, 从而能够缓解资源交互斑块性生境对克隆植物的不利影响.     

Environmentally-induced functional specialization for locally abundant resources in the stoloniferous herb Duchesnea indica

Reciprocally patchy environments, where the availability of two resources are patchily distributed and negatively correlated in each patch, are common in many ecosystems. Interconnected ramets of clonal plants can specialize in the uptake of locally abundant resources. Ramet pairs of the stoloniferous herb Duchesnea indica were grown in reciprocally patchy environments i.e., one ramet of a pair was grown in the high light but low water patch (high light patch) and the other in the low light but high water patch (high water patch). Biomass allocation pattern (root-shoot ratio), morphological traits (leaf area and root length) and physiological traits (photosynthetic rate and chlorophyll content) were altered in a way that potentially enables ramets to enhance the capture of the locally abundant resource (i.e., increase the capture of light resource in the high light patch and of water in the high water patch). As a result,biomass and number of ramets in the connected ramet pairs were greatly improved. Functional specialization of ramets, modified by clonal integration, may have contributed greatly to the growth increase of D. indica in the reciprocally patchy environment.     

资源异质性环境中三种匍匐茎草本植物克隆内资源共享和分株功能特化(英文)

研究了 3种来自中国北方林下、草地和碱化草甸匍匐茎型克隆草本植物绢毛匍匐委陵菜 (PotentillareptansL .var.sericophyllaFranch .)、鹅绒委陵菜 (P .anserinaL .)和金戴戴 (Halerpestesruthenica (Jacq .)Qvcz .)对由高光照低养分斑块和低光照高养分斑块组成的资源交互斑块性生境的适应性对策。当生长于高光照低养分条件下分株(HL分株 )与生长于低光照高养分条件下分株 (LH分株 )之间的匍匐茎连接时 ,3种克隆植物HL分株、LH分株以及整个分株对系统 (HL分株 LH分株 )的生物量均得到显著提高。同时 ,LH分株根冠比显著增加 ,而HL分株根冠比显著下降。这表明 ,当互连分株置于由低光照高养分斑块和高光照低养分斑块组成的异质性环境中时 ,3种植物克隆分株均发生了环境诱导的功能特化。克隆内资源共享以及克隆内不同分株的功能特化有利于整个分株系统对局部丰富资源的获取 ,从而能够缓解资源交互斑块性生境对克隆植物的不利影响     

Effects of light quality and quantity on growth of the clonal plant Eichhornia crassipes

Summary Plant canopy shade reduces photosynthetic photon flux density (PPFD) and ratio of red to far-red light (z). Both effects can cause plants to increase potential for light acquisition through vertical growth and leaf area expansion. Clonal plants such as Eichhornia crassipes might alternatively increase light interception via horizontal growth of stolons or rhizomes and placement of new ramets in less shaded microsites. Effect of simulated canopy shade and component effects of PPFD and z were tested by filtering or adding light uniformly, to a whole group of connected ramets, or locally, to individual ramets within a group. In uniform treatments, low PPFD reduced total growth but low z did not. Low PPFD and low z independently reduced stolon and ramet production and caused etiolation of petioles; effect of low PPFD plus low z on ramet production was greater than that of either factor alone. Lateral clonal growth thus did not seem to be a response to uniform shading; instead, uniformly low PPFD or low z increased partitioning to established ramets. Low z changed partitioning without changing total growth. In local treatments, reduction of growth of individual ramets due to low PPFD and inhibition of new ramet production attributable to spectral composition of light were mitigated when connected ramets were unshaded; plants may respond differently to patchy than to uniform shade.     

Environmentally-induced functional specialization for locally abundant resources in the stoloniferous herb Duchesnea indica

Reciprocally patchy environments, where the availability of two resources are patchily distributed and negatively correlated in each patch, are common in many ecosystems. Interconnected ramets of clonal plants can specialize in the uptake of locally abundant resources. Ramet pairs of the stoloniferous herb Duchesnea indica were grown in reciprocally patchy environments i.e., one ramet of a pair was grown in the high light but low water patch (high light patch) and the other in the low light but high water patch (high water patch). Biomass allocation pattern (root-shoot ratio), morphological traits (leaf area and root length) and physiological traits (photosynthetic rate and chlorophyll content) were altered in a way that potentially enables ramets to enhance the capture of the locally abundant resource (i.e., increase the capture of light resource in the high light patch and of water in the high water patch). As a result,biomass and number of ramets in the connected ramet pairs were greatly improved. Functional specialization of ramets, modified by clonal integration, may have contributed greatly to the growth increase of D. indica in the reciprocally patchy environment.     

Clonal plant allocation to daughter ramets is a simple function of parent size across species and nutrient levels

The evolution of clonal growth is a widespread phenomenon among plant species, characterized by the production of genetically identical clonal fragments (ramets) via rhizomes or stolons that form an interconnected clonal organism (genet). Clonal plant species are known to differ in their investment into ramet production, and exhibit considerable variation in ramet morphology both within and among species. While patterns of resource allocation are thought to be linked to a number of plant characteristics, many analyses are limited by uncertainty in how clonal plants determine the morphology and resources allocated to new ramets. In this study, we attempted to discern what aspects of parent ramets best predicted resource allocation to new daughter ramets, and the relationship between resource allocation and daughter ramet rhizome morphology. We grew two sedge species, Schoenoplectus tabernaemontani and Eleocharis elliptica, in a greenhouse under two levels of fertilizer addition. By harvesting daughter ramets that had initiated stem production, yet remained aphotosynthetic, we were able to isolate parental investment into non-independent daughter ramets at a point where daughter ramet spacer length became fixed. Our results indicate that parent ramets allocated a non-linear proportion of parent rhizome biomass to the production of daughter ramets. Moreover, this relationship was unaffected by environmental nutrient availability. Daughter ramet biomass, in turn, was strongly correlated with daughter ramet spacer length. These observations shed light on key processes governing clonal growth in plants, and their potential application in unifying allocational and morphological perspectives to explore the fitness implications of variability in clonal growth.     

Demographic response to shading and defoliation in two woodland orchids

Many woodland herbs are long-lived, clonal geophytes that have evolved life histories favoring survival over reproduction. We examined the life history responses of natural populations of two woodland orchid species,Cypripedium calceolus andCephalanthera longifolia to defoliation and heavy shading conducted early in the growing seasons of 2002 and 2003. We asked whether, in view of the importance of growth for the survival of geophytes, treated plants were more likely to exhibit reduced flowering than reduced vegetative growth in the seasons following treatment. We also asked whether plants would suffer reduced ramet performance. Both treatments led to significant declines in flower number per ramet, number of leaves per ramet, and mean ramet height relative to controls inCypripedium. However, inCephalanthera, only shaded plants exhibited significant declines in flower number per ramet, and only defoliated plants exhibited declines in mean ramet height. The number of ramets per plant did not decline relative to controls in either species. Thus, these orchids, especiallyCypripedium, appeared to allocate resources preferentially to vegetative growth functions over sexual reproduction. Per-plant variation in leaf and flower number per ramet, as well as in mean ramet height, consistently declined in response to treatment, significantly so in the case of mean ramet height, suggesting that ramets became more similar within genets. These results suggest both similarities and differences in the ways in whichCephalanthera andCypripedium mobilize resources in response to stress.     

Species-specific sprouting pattern in two dioecious Lindera shrubs: The role of physiological integration

Sprouting is recognized as an important genet persistence strategy for clonal woody plants, but the role of sprouting may differ between species and between sexes, depending on physiological integration. We tested the effect of physiological integration on the mortality, recruitment and growth of the sprouting male and female ramets of two closely related dioecious shrubs of Lindera, in a field experiment using girdling manipulation. Although between-sex differences observed were obscure, we found between-species differences in the sprouting patterns. The rates of ramet mortality and recruitment were significantly lower for L. praecox than L. triloba. In L. praecox genets, the ramet production was low, and the main ramets might actively translocate assimilates towards the small sprouted ramets, which then facilitates high ramet growth and survival (sprout-nursing strategy). Meanwhile, in L. triloba genets, although many ramets were recruited, assimilate translocation from the main ramets to the sprouted ramets might be less abundant, which causes high ramet mortality (sprout-turnover strategy). For a more general knowledge of the various sprouting strategies in clonal plants, our study demonstrated that inter-specific comparisons using girdling experiments at the whole-plant level could reveal the role of physiological integration on the link between the sprouting pattern and above-ground structures of clonal plants.