克隆植物对异质生境的适应对策研究进展

生境异质性是自然生态系统的基本特征,植物生长的必需资源和环境胁迫因子均存在着复杂的时间和空间异质性。克隆植物是指在自然条件下具有克隆特性的植物,即可通过与母株相连的芽、根茎、分蘖或枝条等繁殖体产生无性繁殖的植物,这些繁殖体一旦定居便可成为潜在的独立个体。克隆植物具有独特的生境适应策略(如形态可塑性、克隆整合、克隆分工、觅食行为、风险分摊等),面对异质性的生境条件,它可以通过调整自身的生理和形态结构来适应异质生境。目前,对于克隆植物在异质生境适应行为的研究已有很多报道,然而系统性的归纳和总结尚有欠缺。综述了克隆植物在不同资源异质生境(光照、养分、水分)和不同胁迫生境(盐碱胁迫、风沙胁迫、重金属胁迫)下独特的适应对策。最后,针对克隆植物对异质生境的适应对策,进行了总结并对未来的重点研究方向提出建议：(1)时间异质性尺度上的考量;(2)异质性生境中生物因子的调控作用;(3)克隆植物入侵机制;(4)克隆植物在生态修复中的应用潜力。     

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.     

Physiological integration affects growth form and competitive ability in clonal plants

Clonal plants translocate resources through spacers between ramets. Translocation can be advantageous if a plant occurs in heterogeneous environments (division of labour); however, because plants interact locally, any spatial arrangement of ramets generates some heterogeneity in light and nutrients even if there is no external heterogeneity. Thus the capacity of a clonal plant to exploit heterogeneous environment must operate in an environment where heterogeneity is partly shaped by the plant growth itself. Since most experiments use only simple systems of two connected ramets, plant-level effects of translocation are unknown. A spatially explicit simulation model of clonal plant growth, competition and translocation is used to identify whether different patterns of translocation have the potential to affect the growth form of the plant and its competitive ability. The results show that different arrangements of translocation sinks over the spacer system can completely alter clonal morphology. Both runners and clumpers can be generated using the same architectural rules by changing translocation only. The effect of translocation strongly interacts with the architectural rules of the plant growth: plants with ramets staying alive when a spacer is formed are much less sensitive to change in translocation than plants with ramets only at the tip. If translocation cost is low, translocating plants are in most cases better competitors than plants that do not translocate; the difference becomes stronger in more productive environments. Key traits that confer competitive ability are total number of ramet, and their fine-scale aggregation.Co-ordinating editor: J. Tuomi     

养分异质条件下结缕草克隆分株生长、碳水化合物及可溶性蛋白的生理整合

通过对连接和断开的结缕草分株进行不同的养分处理,研究了养分异质条件下结缕草克隆分株生长、碳水化合物及可溶性蛋白含量的变化.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显著提高与其相连子株的地上、地下及总生物量,高养分下分别提高32.5%、22.1%和24.8%,降低根冠比、可溶性糖及非结构性碳水化合物(NSC)含量,高养分下分别降低7.7%、15.2%和13.1%,但是对淀粉、纤维素、可溶性蛋白含量无显著影响.处于中、高养分水平的子株对与其相连的母株生长、碳水化合物及可溶性蛋白含量没有显著影响.养分异质条件下结缕草母株对子株生物量、根冠比、可溶性糖及NSC含量有明显的生理整合,其整合强度与养分梯度呈正比,而对淀粉、纤维素及可溶性蛋白含量没有显现显著的生理整合效应.子株对母株各项指标都没有显著的生理整合,结缕草母株和子株间是一种单向的生理整合.     

不同生境中克隆植物的繁殖倾向

对克隆植物在林下、草原、荒漠、冻原、农田和水域等6种不同类型生境中的繁殖对策及其生态适应意义进行了归纳和总结。对克隆植物有性繁殖与无性繁殖之间的权衡关系及相对重要性随水分、光照和温度的变化规律进行了阐述。分析表明,在环境条件严酷地区,克隆植物的无性繁殖在种群更新中的相对重要性有所增加。对两种繁殖方式的生态功能特性进行比较,综述了有关二者权衡关系的研究成果及影响二者权衡关系的部分因素。从研究手段上对克隆植物繁殖对策的研究进行了思考与展望。     

No evidence for nutrient foraging in root-sprouting clonal plants

Clonality is defined as vegetative reproduction via the production of ramets, which are, at least initially, connected by spacers. In general, there are three types of spacers of two origins. Whereas stolons are aboveground spacers, rhizomes are belowground spacers; however, both of stem origin. The third type of spacers are roots in root-sprouting plants. The possibility of foraging in clonal plants has attracted broad interest among ecologists but has been experimentally documented only for stoloniferous clonal plants foraging for light. Foraging for belowground resources has yet to be demonstrated, perhaps because tests of foraging have focused on clonal plants that spread laterally via stolons or rhizomes, i.e. stem organs. Lateral spread based on sprouting roots has not been considered even though, in addition to functioning as conduits between ramets, root spacers are able to sense and take up nutrients. We therefore hypothesized that root-sprouting clonal plants may be able to directly react to environmental heterogeneity and exhibit nutrient foraging. To test this hypothesis, we conducted two experiments with root-sprouters in nutrient-heterogeneous and -homogeneous environments. We found that plants produced more biomass when growing in a heterogeneous environment than in a homogeneous environment and that root biomass was greater in the nutrient-rich patches than in nutrient-poor patches. However, the number of ramets did not differ between patches in the heterogeneous environment. We conclude that plants whose clonality is based on roots, similarly as plants whose clonality is based on stolons or rhizomes, do not exhibit accumulation of ramets in nutrient-rich patches. Foraging at the organ level, i.e. by roots, seems to be more probable in this clonal group. To analyse how clonal plants with different clonal strategies perceive and react to environmental heterogeneity, researchers must account for the high variability in clonal growth forms and in scales of environmental heterogeneity.     

不同水盐生境下芦苇湿地植被及土壤碳氮磷生态化学计量特征

在黄河三角洲选取3种典型的芦苇湿地群落为对象,即:故道区(1996年改道的黄河故道河岸带芦苇群落)、新生区(现行黄河河岸带新生芦苇群落)和潮水区(远离新旧河道但受潮汐影响的潮滩芦苇群落),研究黄河改道对3种生境芦苇各器官(茎、叶、根状茎、须根)和土壤剖面碳(C)、氮(N)、磷(P)含量变化及化学计量特征的影响.结果 表...     

Patchy habitats, division of labour and growth dividends in clonal plants

Natural habitats are patchy in quality. in clonal plants, resource-acquiring structures often occupy sites that differ in quality. Clonal plants can display division of labour in resource-acquisition duties, manifested as local specialization by ramets, which enhances acquisition of each resource from sites of greatest abundance. Physiological integration can re-distribute resources internally from sites of acquisition to clone parts sited where the same resources are scarce. Recent research is showing that such specialization and resource sharing is a highly efficient strategy for acquiring resources and that it can result in considerably greater growth when resources are heterogeneously distributed than when the same quantity of resources is distributed homogeneously.     

Physical connection decreases benefits of clonal integration in Alternanthera philoxeroides under three warming scenarios

Physical connection between ramets usually allows clonal plants to perform better but can have the opposite effects in some cases. Clonal integration and the effects of climate warming have been extensively studied, but to date little is known about how climate warming affects the benefits of clonal integration. We conducted a field experiment in which Alternanthera philoxeroides segments with connected and severed stolons were subject to four climate regimes (ambient, day warming, night warming and daily warming), and measured final biomass, number of ramets and total length of stolons. Across the three warming treatments, temperature rise suppressed growth of clonal fragments with connected stolons but increased growth of fragments with severed stolons; temperature rise affected the biomass of distal ramets but not proximal ramets, and had similar effects on the numbers of proximal and distal ramets. When the three warming treatments were considered separately, they had contrasting consequences for the benefits of clonal integration. Specifically, when fragments were exposed to day and night warming, physical connection evened out the advantages of clonal integration that occur under ambient conditions; when fragments were exposed to daily warming, physical connection led to smaller clonal plants. These findings suggest that physical connection between ramets may be disadvantageous to overall performance of A. philoxeroides fragments under climate warming, and also indicate that the net consequences of daily warming outweigh those of day or night warming.     

Effects of clonal integration on allelopathy of invasive plant Wedelia trilobata under heterogeneous light conditions

异质光照条件下克隆整合对入侵植物南美蟛蜞菊化感作用的影响 植物入侵已成为全球生态系统最严重的威胁之一。当具有克隆生长能力的入侵植物入侵或定殖到新的生境时,它们相互连接的分株可能受到异质光照的影响。在异质光照条件下,克隆整合对入侵植物化感作用的影响尚不清楚。为研究异质光照条件下克隆整合对入侵植物南美蟛蜞菊(Wedelia trilobata) 化感作用的影响,采用两个连续分株的克隆片段进行了盆栽试验。较老的分株暴露在全光下,而年轻的分株则受到20%的全光照。同时,每个克隆片段的年轻分株与目标植株(一个番茄苗)在盆栽中相邻生长。南美蟛蜞菊的两个连续分株之间的匍匐茎设置切断和不切断两种处理。另外,两株番茄幼苗 (一株作为目标植株)在盆栽中相邻生长作为对照。研究结果表明,当与目标植物相邻生长的南美蟛蜞菊分株之间的匍匐茎保持完整时,目标植株的生物量积累、叶片叶绿素和氮含量、叶绿素荧光参数和净光合速率及其根长和活性相较于匍匐茎切断处理显著降低。异质光照条件下连续两个分株之间碳水化合物的运输或共享可以增强20%全光处理下年轻分株的化感作用。克隆整合在异质光照条件下对具有克隆生长能力的入侵植物的入侵或定殖具有重要意义。     

Physiological integration in the clonal perennial herb Trifolium repens L.

Summary Translocation of ¹⁴C-labelled carbohydrates between the parent stolon and branches, and among branches, of Trifolium repens plants was investigated in two glasshouse experiments to determine patterns of physiological organisation in this clonal species. Differential defoliation treatments were applied to the parent stolon and/or branches to test the sensitivity of translocation to the short-term carbon needs of defoliated sinks. Strong reciprocal exchange of carbohydrate between the parent stolon and branches was observed, with 18 41% of the ¹⁴C exported from leaves on the parent stolon moving to branches, while branches simulta-neously exported 25% (for old source branches) to 54% (for young source branches) of the ¹⁴C they assimilated to the parent plant, including translocation to other branches. Branch-to-branch translocation occurred both acropetally and basipetally. Parent-to-branch, branch-to-parent and branch-to-branch carbon fluxes all increased in response to defoliation of the sink, at the expense of carbon supply to stolon tissue or roots of the source module. Reduced export to stolon tissue of the parent axis played a major role in facilitating C reallocation from leaves on the parent stolon to defoliated branches. The observed patterns of C allocation and translocation could be adequately explained by accepted source-sink theory, and are consistent with a high degree of intra-plant physiological integration in resource supply and utilisation. This information provides mechanistic explanations for aspects of the growth dynamics and ecological interactions of T. repens in the patchy environment of a grazed pasture.     

Nitric oxide is involved in integration of UV‐B absorbing compounds among parts of clonal plants under a heterogeneous UV‐B environment

In nature, ultraviolet‐B (UV‐B) radiation is highly heterogeneous, both spatially and temporally. Plants exposed to UV‐B radiation produce UV‐B absorbing compounds that function as a protective filter. For clonal plants under heterogeneous UV‐B radiation conditions, integration among ramets can allow irradiated ramets to benefit un‐irradiated ramets by causing them to increase their UV‐B absorbing compounds content. In this study, we evaluated integration between pairs of clonal ramets of Glechoma longituba under heterogeneous or homogeneous UV‐B conditions. We determined the levels of UV‐B absorbing compounds, nitric oxide (NO) and hydrogen peroxide (H₂O₂) and measured the activity of phenylalanine ammonia‐lyase (PAL) in connected ramet pairs under homogeneous or heterogeneous UV‐B conditions. Under heterogeneous UV‐B conditions, the UV‐B absorbing compounds content increased in leaves of irradiated and un‐irradiated ramets, but not in the connecting stolons. The NO content increased in irradiated and un‐irradiated leaves and stolons, but the H₂O₂ content did not. Application of NO synthesis inhibitors and an NO blocker to irradiated ramets blocked the increase in UV‐B absorbing compounds and PAL activity in un‐irradiated ramets. These results suggested that NO is involved in the integration process for UV‐B absorbing compounds among ramets. Our findings suggested that a UV‐B‐induced increase in NO transmits a signal to un‐irradiated ramets via the stolon, leading to an increase in PAL activity and UV‐B absorbing compounds content. The internal translocation of signal enables members of clonal networks to function as a whole unit and to mount an efficient defensive response to localized UV‐B radiation.     

Physiological and biochemical changes in plants under waterlogging

Waterlogging usually results from overuse and/or poor management of irrigation water and is a serious constraint with damaging effects. The rapidly depleting oxygen from submerged root zone is sensed and plant adjusts expressing anaerobic proteins. Plant cells shift their metabolism towards low energy yielding anaerobic fermentation pathways in the absence of oxygen. Structural modifications are also induced as aerenchyma formation and adventitious rootings, etc. Studies at molecular and biochemical levels to facilitate early perception and subsequent responses have also been worked out to produce resistant transgenic plants. This review explores the sequential changes of plant responses at different levels regarding their defense strategies and efforts made to enhance them, tailoring crucial regulators so that they can withstand waterlogging stress.     

Bicarbonate use in three aquatic plants

Our study aimed to test the ability of aquatic plants to use bicarbonate when acclimated to three different bicarbonate concentrations. To this end, we performed experiments with the three species Ceratophyllum demersum, Egeria densa, Lagarosiphon major to determine photosynthetic rates under varying bicarbonate concentrations. We measured bicarbonate use efficiency, photosynthetic performance and respiration. For all species, our results revealed that photosynthetic rates were highest in replicates grown at low alkalinity. Thus, E. densa had approx. five times higher rates at low (264 ± 15 μmol O₂ g⁻¹ DW h⁻¹) than at high alkalinity (50 ± 27 μmol O₂ g⁻¹ DW h⁻¹), C. demersum had three times higher rates (336 ± 95 and 120 ± 31 μmol O₂ g⁻¹ DW h⁻¹), and L. major doubled its rates at low alkalinity (634 ± 114 and 322 ± 119 μmol O₂ g⁻¹ DW h⁻¹). Similar results were obtained for bicarbonate use efficiency by E. densa (136 ± 44 and 43 ± 10 μmol O₂ mequiv. L⁻¹ g⁻¹ DW h⁻¹) and L. major (244 ± 29 and 82 ± 24 μmol O₂ mequiv. L⁻¹ g⁻¹ DW h⁻¹). As to C. demersum, efficiency was high but unaffected by alkalinity, indicating high adaptation ability to varied alkalinities. A pH drift experiment supported these results. Overall, our results suggest that the three globally widespread worldwide species of our study adapt to low inorganic carbon availability by increasing their efficiency of bicarbonate use.     

Physiological integration ameliorates effects of serpentine soils in the clonal herb Fragaria vesca

Small-scale heterogeneity in soil characteristics and the facility of clonal systems to spread may lead to situations where parent ramets in favourable microhabitats are connected to offspring in stressful conditions. Clonal plants are physiologically integrated if connections among ramets allow transport of resources. Thus, ramets in favourable habitats may provide support to developing or stressed ramets. We examined effects of integration in Fragaria vesca growing in patches of contrasting quality (potting compost vs serpentine soil). Serpentine soil was used to create unfavourable growing conditions. We assessed whether survival, biomass and photosynthetic efficiency (estimated by fluorescence and reflectance) of parents and offspring were affected by integration and soil quality. Integration increased photochemical efficiencies of parents but more consistently in parents with offspring growing in serpentine soils. We suggest that the assimilate demand from offspring enhanced the photosynthetic efficiency of parents by a mechanism of feedback regulation. This result extends the concept of physiological integration in clonal plants to include photochemical responses. Connected parents also showed significantly higher biomass than disconnected parents. In our system, integration did not entail costs for the whole clone in terms of biomass. Integration also improves the survival, growth and photochemical efficiency of developing ramets, suggesting that integration represents a mechanism for increasing survival in stressful habitats, as the serpentine soils.     

Benefits of clonal integration between interconnected ramets of Vallisneria spiralis in heterogeneous light environments

Interconnected ramets of the submersed macrophyte Vallisneria spiralis were subjected to two homogeneous treatments (shading or not shading whole plants) and two heterogeneous treatments (only shading basal or apical ramets of plants). The benefits and costs of clonal integration between connected ramets grown in heterogeneous treatments were examined. Results showed that shading apical ramets induced significant benefits to the performance of whole plant in terms of dry weight per plant (P < 0.01) and number of ramets per plant (P < 0.05). Especially for the unshaded basal ramets, their dry weight, number of ramets, number of branches and total stolon length were 89%, 30%, 29% and 58% higher than the corresponding ramets in homogeneous treatment, respectively. Compared to their controls in homogeneous treatments, unshaded basal ramets produced more leaf mass (0.15 g versus 0.11 g) whereas shaded apical ramets produced more root mass (0.012 g versus 0.008 g). However, there was a different pattern of integration when basal ramets were shaded. Shading basal ramets led to a significant decrease in stolon growth, but the individual performance of shaded ramets improved. Cost-benefit analyses revealed that dry weight per ramet of basal shaded ramets was 31% greater than that of basal shaded ramets in the homogeneous treatment. We can conclude that V. spiralis can benefit from clonal integration in heterogeneous light environments, but that the scale of these benefits is related to the quality of light environments where the clone become established.     

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。     

Bats and aquatic habitats: a review of habitat use and anthropogenic impacts

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异质性光照下匍匐茎草本狗牙根克隆整合的耗益

在一盆栽实验中,将匍匐茎草本植物狗牙根克隆分株对进行异质性光照处理,对远端分株(幼年)分别进行不同程度的遮荫,并对分株对之间的匍匐茎进行保持连接或是切断处理。结果显示:在中度遮荫时,切断匍匐茎连接显著降低了远端分株的生物量和净光合速率(Pn);在重度遮荫时,切断匍匐茎连接显著降低了远端分株的分株数、叶片数、生物量、净光合速率(Pn)、最大光量子产量(Fv/Fm)和实际光量子产量(ФPSⅡ),克隆整合缓解了遮荫胁迫对远端分株生长的影响。在中度遮荫时,保持匍匐茎连接显著提高了近端分株的净光合速率(Pn);在重度遮荫时,保持匍匐茎连接显著提高了近端分株的最大光量子产量(Fv/Fm)、实际光量子产量(ФPSⅡ)和净光合速率(Pn),克隆整合使得相连未受胁迫近端分株的光合效率表现出补偿性增加。相比于匍匐茎切断处理,当远端分株遭受重度遮荫胁迫时,克隆整合引起近端未受胁迫分株生物量显著降低,而整个克隆片段生物量没有显著变化。因此,当远端分株遭受重度遮荫胁迫时,匍匐茎草本狗牙根可能采取一种风险分摊策略以降低基株死亡风险,这样一种策略对于维持克隆植物在异质性生境中基株适合度具有重要意义。