Herbivores Influence the Growth,Reproduction, and Morphology of a Widespread Arctic Willow

Shrubs have expanded in Arctic ecosystems over the past century, resulting in significant changes to albedo, ecosystem function, and plant community composition. Willow and rock ptarmigan (Lagopus lagopus, L. muta) and moose (Alces alces) extensively browse Arctic shrubs, and may influence their architecture, growth, and reproduction. Furthermore, these herbivores may alter forage plants in such a way as to increase the quantity and accessibility of their own food source. We estimated the effect of winter browsing by ptarmigan and moose on an abundant, early-successional willow (Salix alaxensis) in northern Alaska by comparing browsed to unbrowsed branches. Ptarmigan browsed 82–89% of willows and removed 30–39% of buds, depending on study area and year. Moose browsed 17–44% of willows and browsed 39–55% of shoots. Browsing inhibited apical dominance and activated axillary and adventitious buds to produce new vegetative shoots. Ptarmigan- and moose-browsed willow branches produced twice the volume of shoot growth but significantly fewer catkins the following summer compared with unbrowsed willow branches. Shoots on browsed willows were larger and produced 40–60% more buds compared to unbrowsed shoots. This process of shoot production at basal parts of the branch is the mechanism by which willows develop a highly complex “broomed” architecture after several years of browsing. Broomed willows were shorter and more likely to be re-browsed by ptarmigan, but not moose. Ptarmigan likely benefit from the greater quantity and accessibility of buds on previously browsed willows and may increase the carrying capacity of their own habitat. Despite the observed tolerance of willows to browsing, their vertical growth and reproduction were strongly inhibited by moose and ptarmigan. Browsing by these herbivores therefore needs to be considered in future models of shrub expansion in the Arctic.     

Spatio-temporal patterns of ptarmigan occupancy relative to shrub cover in the Arctic

Rock and willow ptarmigan are abundant herbivores that require shrub habitats in arctic and alpine areas. Shrub expansion is likely to increase winter habitat availability for ptarmigan, which in turn influence shrub architecture and growth through browsing. Despite their ecological role in the Arctic, the distribution and movement patterns of ptarmigan are not well known, particularly in northern Alaska where shrub expansion is occurring. We used multi-season occupancy models to test whether ptarmigan occupancy varied within and among years, and the degree to which colonization and extinction probabilities were related to shrub cover and latitude. Aerial surveys were conducted from March to May in 2011 and April to May 2012 in a 21,230 km² area in northeastern Alaska. In areas with at least 30 % shrub cover, the probability of colonization by ptarmigan was >0.90, indicating that moderate to extensive patches of shrubs (typically associated with riparian areas) had a high probability of becoming occupied by ptarmigan. Occupancy increased throughout the spring in both years, providing evidence that ptarmigan migrated from southern wintering areas to breeding areas north of the Brooks Range. Occupancy was higher in the moderate snow year than the high snow year, and this was likely due to higher shrub cover in the moderate snow year. Ptarmigan distribution and migration in the Arctic are linked to expanding shrub communities on a wide geographic scale, and these relationships may be shaping ptarmigan population dynamics, as well as rates and patterns of shrub expansion.     

松嫩平原野古草种群构件结构动态

野古草是根茎型无性系禾草,在松嫩平原草甸经常形成单优种群落。采用单位面积挖掘取样、分株按营养繁殖世代划分龄级、根茎按实际生活年限划分龄级的方法,对松嫩平原单优群落和混生群落的野古草种群构件结构进行了调查与分析。结果表明,在生长季初期两群落野古草种群均以春性分株和根茎芽占优势,且分株及芽构件结构相对稳定,芽库的输出率单优群落为80.4%,混生群落为62.5%;整个生长季分株由2—3个龄级组成,1a分株数量是2a的2.9—10.2倍,其生物量各月份所占比例平均为93%,随着龄级的增加依次明显减少,呈增长型年龄结构;根茎由3—4个龄级组成,根茎累积长度及生物量均以2a占绝对优势,为稳定型年龄结构;分株生产力1a明显高于2a,对种群贡献最大;根茎贮藏力除个别月份以3a、4a最高外,两群落大部分以2a最高,在生长季后期,1a根茎物质积累的速率最快。     

Position of tillers in a clone determines their ontogeny: example of the clonal grass <Emphasis Type="Italic">Phalaris arundinacea</Emphasis>

For practical reasons, all ramets of a clonal plant are considered to be independent individuals in demographical studies, but there are reports that ramet ontogeny is affected by their position in a clone. This means that ramets growing from buds located in certain parts of the rhizome system differ in size and probability of flowering from ramets emerging from buds at various positions. We studied the growth characteristics and ontogeny of tillers (ramets) of a rhizomatous grass in relation to their position in a clone and examined whether it is affected by the disturbance regime. We assessed the position of tillers in a clone, the developmental stage of their apical meristem and growth characteristics in mown and unmown stands of the rhizomatous grass Phalaris arundinacea at two sites, one representing optimal (lowland) and the other suboptimal (highland) environmental conditions. We found that tiller characteristics and the probability of a tiller to start flowering are related to the tiller position in a clone. Tiller differences were preserved even in suboptimal environmental conditions but disappeared after mowing, which equalized size and ontogeny. Our results imply that ramets of clonal plants are not independent demographical units, thus their position in a clone should be taken into account.     

Growth of coyote willow and the attack and survival of a mid-rib galling sawfly,Euura sp.

We studied the relationship between variation in age and shoot characteristics of the host plant Salix exigua Nuttall (coyote or sandbar willow) and the attack and survival of Euura sp. (an unnamed leaf-midrib galling sawfly). Variation in shoot characteristics resulted from reduced growth as willow ramets aged. Mean shoot length per ramet and mean longest leaf length per shoot decreased by 95% and 50% respectively between 1- and 9-year-old willow ramets. All measured shoot characteristics-shoot length, longest leaf length, number of leaves per shoot, and mean internode length-were significantly negatively correlated with ramet age (r ² ranged from –0.23 to –0.41). Correlations between shoot characteristics were highly positive, indicating that plants also grew in a strongly integrated fashion (r ² ranged from 0.54 to 0.85). Four hypotheses were examined to explain sawfly attack patterns. The host-plant hypothesis was supported in explaining enhanced larval sawfly survival through reduced plant resistance. As willow ramets aged, the probability of Euura sp. attack decreased over 10-fold, from 0.315 on 1-year-old ramets to 0.024 on 2- to 9-year-old ramets. As shoot length increased, the probability of sawfly attack increased over 100-fold, from 0.007 on shoots <100 mm, to 0.800 on shoots in the 1001–1100 mm shoot length class. These attack patterns occurred even though 1-year-old ramets and shoots >500 mm each represented less than 2% of the total shoots available for oviposition. Host plant induced mortality of the egg/early instar stage decreased by 50% on longer leaves and was the most important factor determining survival differences between vigorous and non-vigorous hosts. Sawfly attack was not determined by the resource distribution hypothesis. Although shoots <200 mm contained 82% of the total leaves available, they contained only 43% of the galls initiated. The attack pattern also was not explained by the gall volume hypothesis. Although gall volume increased on longer shoots, there was no significant variation in mid or late instar mortality over shoot length, as would be expected if food resources within smaller galls were limited. The natural enemy attack hypothesis could not explain the pattern of oviposition since predation was greater on longer shoots and leaves. In addition, larval survival was related to oviposition behavior. Due to a 69% reduction in late instar death and an 83% reduction in parasitism, survival of progeny in galls initiated close to the petiole base was 2.8 times greater than in galls initiated near the leaf tip. A 75% reduction in gall volume over this range of gall positions may account for the observed increases in late instar mortality and parasitism.     

松嫩平原全叶马兰种群无性系分株的建成及其年龄结构

全叶马兰（Kalimeris integrifolia Turcz.)为根蘖型多年生草本菊种植物。在生长季后期,新生肉质细根向地表生长形成根蘖芽,根蘖芽成株后形成主根。经过对大量样本观察。创建了按形态特征对根蘖型草本植物种群无性系分株和芽的年龄划分方法,以及对主根生活年限的估计方法。分株年龄是按主根芽再生的次数划分,新根蘖芽成株为1a,1a分株死亡后从主根再萌生的分株为2a,2a分株死亡后从主根再萌生的分株为3a。新根蘖芽为0a芽,其它龄级与分株的龄级相一致,即1a分株主根芽为1a芽,2a分株主根芽为2a芽,在松嫩平原割草草场,全叶马兰种群由3个龄级无性系分株组成,呈稳定型年龄结构。分株主根的生活年限最多为4年,分株的生产力种群水平以2a最高,个体水平以1a最低,并随着龄级的增加呈增长趋势,其中,在生长季初期以3a分株的生产力最高,进入中期以后则1a最低,并随着龄级的增加呈增长趋势。其中,在生长季初期以3a分株的生产力最高,进入中期以后则1a分株的相对生长速率最大,用秋末新生根蘖数量和现时种群数量可以较准确预报翌年种群数量和年龄结构。芽库中潜在种群数量是现时种群数量的2倍以上,但不意味着种群将发生疯长,而是为替补所做的充分准备。     

Density-dependent regulation of ramet recruitment by the red:far-red ratio of solar radiation: a field evaluation with the bunchgrass Schizachyrium scoparium

Depressions in the red to far-red ratio (R:FR) of solar radiation arising from the selective absorption of R (600–700 nm) and scattering of FR (700–800 nm) by chlorophyll within plant canopies may function as an environmental signal directly regulating axillary bud growth and subsequent ramet recruitment in clonal plants. We tested this hypothesis in the field within a single cohort of parental ramets in established clones of the perennial bunchgrass, Schizachyrium scoparium. The R:FR was modified near leaf sheaths and axillary buds at the bases of individual ramets throughout the photoperiod without increasing photosynthetic photon flux density (PPFD) by either (1) supplementing R beneath canopies to raise the naturally low R:FR or (2) supplementing FR beneath partially defoliated canopies to suppress the natural R:FR increase following defoliation. Treatment responses were assessed by simultaneously monitoring ramet recruitment, PPFD and the R:FR beneath individual clone canopies at biweekly intervals over a 12-week period. Neither supplemental R nor FR influenced the rate or magnitude of ramet recruitment despite the occurrence of ramet recruitment in all experimental clones. In contrast, defoliation with or without supplemental FR beneath clone canopies reduced ramet recruitment 88% by the end of the experiment. The hypothesis stating that the R:FR signal directly regulates ramet recruitment is further weakened by evidence demonstrating that (1) the low R:FR-induced suppression of ramet recruitment is only one component of several architectural modifications exhibited by ramets in response to the R:FR signal (2) immature leaf blades, rather than leaf sheaths or buds, function as sites of R:FR perception on individual ramets, and (3) increases in the R:FR at clone bases following partial canopy removal are relatively transient and do not override the associated constraints on ramet recruitment resulting from defoliation. A depressed R:FR is probably of greater ecological significance as a signal of competition for light in vegetation canopies than as a density-dependent signal which directly regulates bud growth and ramet recruitment.     

High levels of inter-ramet water translocation in two rhizomatous Carex species,as quantified by deuterium labelling

We studied water trnaslocation between interconnected mother and daughter ramets in two rhizomatous Carex species, using a newly developed quantitative method based on deuterium tracing. Under homogeneous conditions, in which both ramets were subjected either to wet or dry soil, little water was exchanged between the ramets. When the ramet pair was exposed to a heterogeneous water supply, water translocation became unidirectional and strongly increased to a level at which 30–60% of the water acquired by the wet ramet was exported towards the dry ramet. The quantity of water translocated was unrelated to the difference in water potential between the ramets, but highly correlated to the difference in leaf area. In both species, the transpiration of the entire plant was similar under heterogeneous and homogeneous wet conditions. This was a direct result of an increase in water uptake by the wet ramet in response to the dry conditions experienced by the interconnected ramet. In C. hirta, the costs and benefits of integration in terms of ramet biomass paralleled the responses of water consumption. This species achieved a similar whole-plant biomass in heterogeneous and homogeneous wet treatments, and water translocation was equally effective in the acropetal and basipetal directions. In C. flacca, responses of biomass and water consumption did not match and, under some conditions, water translocation imposed costs rather than benefits to the plants of this species. It is concluded that enhanced resource acquisition by donor ramets may be of critical importance for the net benefits of physiological integration in clonal plants.     

Clonal integration and its ecological significance in Hedysarum laeve, a rhizomatous shrub in Mu Us Sandland

Hedysarum laeve, a rhizomatous clonal half-shrub, commonly dominates in inland dunes in semiarid areas of northern China. This species propagates vegetatively by the extension of horizontal rhizomes resulting in programmed reiteration of apical and/or axillary meristems. In this study, ¹⁴C labeling and experimental defoliation were employed to test the photosynthate translocation within the interconnected parent–daughter ramet pairs. A proportion of ¹⁴C-photosynthates was transported from the parent ramet into the daughter ramet, the roots of the daughter ramet, and the rhizome; these three components showed more than 70% sink activity after 24-h translocation. On the other hand, the basipetal translocation (from daughter ramet into parent ramet) was relatively small with sink activity of less than 5%, but sink activity of the rhizome exceeded 10%. Defoliation had an influence on the photosynthate translocation between parent and daughter ramets. The intact parent ramets significantly increased their ¹⁴C-photosynthate translocation into defoliated daughter ramets when compared to intact daughter ramets. The daughter ramets transported significantly more ¹⁴C-photosynthates to the defoliated parent ramets than to the intact parent ramets. A portion of ¹⁴C-photosynthates was transported into the rhizome from both parent and daughter ramets, indicating that the rhizome is supported by both ramets for photosynthates. The clonal integration between ramets of the species through rhizome connection may confer benefit both to the ramets and the genet in adverse environments. Received: April 12, 2001 / Accepted: November 26, 2001     

Reproductive demography of ramets and genets in a rhizomatous clonal plant <Emphasis Type="Italic">Convallaria keiskei</Emphasis>

Clonal growth occurring below the ground makes it difficult to identify individuals and demonstrate the demographic features of a focal plant species. In this study, genotypically identified ramets of a rhizomatous clonal herb, Convallaria keiskei Miq., were monitored for their growth, survival, and reproduction from 2003 to 2006. After the monitoring period, their subterranean organs were excavated to explore the underground connections of established ramets and the direction of clonal growth. We then combined data on the fate of the monitored ramets with the information of rhizome connections, clarifying reproductive demography at both the ramet and genet levels. Although each ramet initiated both sexual reproduction (via flowering) and clonal growth, clonal growth tended to precede sexual reproduction. In a surveyed genet, 51.0% of ramets produced flowers and 29.6% generated clonal offspring during the study period. Consequently, we clarified the reproductive demography of C. keiskei: clonal growth tended to precede flowering in a ramet, and a genet can keep reproducing every season at the genet level, despite a ramet not having inflorescence every year.     

Plasticity in bud demography of a rhizomatous clonal plantLeymus chinensis L. in response to soil water status

We surveyed the bud demography ofLeymus chinensis L plants along a soil-moisture gradient that was caused by a flood in 1998 on the Song-nen Plain in northeastern China. The number of vegetative buds per ramet was influenced by soil water content, with regression curves being quadratic and the opening of the parabola pointing downward. In addition, the optimum regression models for the numbers of rhizomatous buds and tiller buds relative to soil water resulted in a quadratic parabola and exponential curve, respectively. Vegetative buds flourished between August and October, with plants producing more of those buds on flooded plots than on control sites. The number of rhizomatous buds per ramet was much higher than for tiller buds throughout most of the growing season, and production of the former was more apt to be affected by soil water status. This observed superiority of rhizomatous bud production was thought to be a consequence of the whole-plant adjustment that was stimulated by an abnormally high moisture content. It could also be interpreted as a strategy for “escape” from disadvantageous overly wet conditions. Moreover, the position-based preference for bud emergence along the ramets could be an underlying mechanism for selective ramet placement.     

扎龙湿地不同生境芦苇分株的形态特征及生物量分配的异速关系

不同环境条件下的植物个体可以表现出形态特征的分异和物质分配的权衡与调整。采用大样本抽样调查与统计分析方法,比较研究扎龙湿地不同生境芦苇(Phragmites Australis)生殖株和营养株的形态特征以及生物量分配的异速关系。结果表明：在9月末,盐碱生境、旱生生境、湿生生境和水生生境芦苇分株的生长表现出较大的生态可塑性,株高和株重均以盐碱生境最小,水生生境最大,最大值与最小值的比值分别为1.3—3.3和1.8—5.1,分株生长在种群间的变异度高于种群内,与营养株相比,生殖株的变异度较低;分株的支持分配与生产分配的比值为1.8—4.2,生产分配以盐碱生境最高,以水生生境最低,而支持分配和生殖分配表现与生产分配相反的序位;生殖株的花序长和花序重与株高间呈直线函数形式增长,株高和株重低于种群平均值的20%和35%的分株不进行有性生殖;叶重、叶鞘和茎重以及分株重与株高间呈幂函数形式的异速生长关系。植物通过改变个体的形态特征以及调整构件间生物量分配适应不同环境,而受遗传因素控制的构件间生长关系却相对稳定。     

Allocation of resources within mountain birch canopy after simulated winter browsing

As a response to browsing, birches are known to produce fewer but larger, more nutritious leaves, with enhanced palatability for herbivores. We simulated winter browsing in ramets of mountain birch ( Betula pubescens ssp. czerepanovii ) to find out whether it decreases subsequent foliage biomass and alters the number and type of shoots. After removal of a considerable proportion of buds (up to 35%) in late winter, the birches were able to compensate for the lost leaf biomass in the following summer; there were no differences in total leaf biomass between winter-clipped and control ramets. This indicates that foliage growth was limited by the total amount of stored resources, not by the number of buds. Depending on the position of the buds removed, different mechanisms were responsible for the compensation. After removal of apical buds, the number of leaves decreased significantly but leaves were larger than in control ramets. Removal of the same mass of basal buds – containing similar amount of carbohydrates and proteins as in the treatment removing apical buds – activated dormant buds, especially in apical locations, so that leaf number was similar as in the controls; consequently, size of individual leaves increased only slightly. Thus, while the total leaf biomass in a tree seems to be limited by resources from source organs, the distribution of resources among different canopy sections is controlled by their relative sink strengths. In terms of leaf biomass, apical parts are able to compensate for bud loss by increasing shoot number, basal parts only by increasing leaf size.     

慈竹构件和分株水平总黄酮含量的变化

黄酮类化合物是植物体内重要的次生代谢物质之一。本文采用AlCl3 比色法测定了慈竹构件和分株水平总黄酮含量的变化 ,以探讨其生理生态适应性。结果表明 ,在地上 3个构件中 ,叶的总黄酮含量显著大于枝和秆 ;1龄分株叶的总黄酮含量显著大于其它 4个龄级 ,而枝和秆的含量在 5个年龄间无显著差异 ;分株上层叶和枝条的总黄酮含量显著多于中下两层的相应构件。叶龄对慈竹叶总黄酮含量有显著的影响 ,1龄、 7龄和 9龄叶片的总黄酮含量明显高于其它 6个龄级 ,但是叶龄和总黄酮含量之间无显著线性相关。同时 ,叶总黄酮含量对季节的变化敏感 ,2、 4和 12月含量最高 ,5月和 8月含量最低。在分株水平 ,1龄分株的总黄酮含量显著大于其它 4个龄级。这些结果说明慈竹的总黄酮含量对环境变化具有可塑性 ,分株水平与叶构件水平总黄酮含量对分株年龄的反应格恐相同。     

Evolutionary impacts of the goldenrod ball gallmaker on Solidago altissima clones

Summary Three ramet clones of Solidago altissima were grown under greenhouse conditions to determine the effects of varying levels of attack by the goldenrod ball gallmaker (Eurosta solidaginis) on biomass allocation, leaf senescence rate and rhizome connections among ramets. The results, examined at both the individual ramet level and the clone level, showed that galled ramets became isolated from their clone through deterioration of rhizome connections. Gall effects were only observed at the ramet level although rhizome connection effects were detected at both the ramet and clone levels. The goldenrod ball gallmaker may therefore have little evolutionary impact on large clones but could appreciably affect newly established clones.     

Ramet dynamics in a centrifugally expanding clonal sedge: a matrix analysis

Carex humilis is a clonal sedge that can form distinct rings of densely aggregated ramets. We hypothesize that rings form because both production of new ramets and ramet dispersal are positively correlated to ramet size. This would lead to an overrepresentation of fast-moving and large ramets with high ramet production at the periphery, whereas slow-moving and small ramets with low ramet production would mainly be found in the interior of rings. We use matrix models to analyse how ramet populations both at the periphery and in the interior develop in the absence of ramet dispersal. We found that the stable size class distributions of ramets predicted by the models were not different from the distributions found in the field. Also, the asymptotic ramet population growth rates (λ₁) were the same. Hence, we conclude that rings would form even in the absence of a link between ramet dispersal and ramet production. Further analysis of the matrix models showed that the ramet population increases at the periphery but decreases in the interior of rings because medium and large ramets produce fewer large ramets in the interior than at the periphery. We also found that the temporal variance in λ₁ and transitions rates during the four study years was much higher at the periphery than in the interior. Our results suggest that rings may form because C. humilis ramets use below-ground resources from a much larger area than the one covered by the shoots. As the clone grows larger, the soil volume available to the ramets in the interior decreases because their access to soil outside the ring is cut-off by the ramets at the periphery. Ramet density in the interior is therefore decreasing.     

Metallothionein levels in willow ptarmigan (<Emphasis Type="Italic">Lagopus lagopus</Emphasis>) populations with different natural loads of cadmium

The tissues of willow ptarmigan in some Norwegian mountain areas contain elevated concentrations of cadmium (Cd). It is not known whether such high Cd levels would have negative impacts in otherwise healthy populations of this species. The aim of the current study was to clarify relationships between hepatic and renal metallothionein (MT) and Cd concentrations in willow ptarmigan to assess effects from this metal. The study reported here was undertaken on willow ptarmigan from the Kongsvoll area, with a naturally high Cd load, and the Essand area, with a naturally low Cd load. Cd values in liver and kidney in willow ptarmigan from Kongsvoll were significantly higher than in willow ptarmigan from Essand. The MT content in both tissues was also highest in willow ptarmigan from Kongsvoll. The MT concentration in kidney was twice that in liver for ptarmigan from both areas and at all times of the year. The MT level in both liver and kidney varied greatly throughout the season, with the highest content in spring (May). The variation was greatest in liver. The total material showed a significant linear relationship between Cd and MT levels in both liver and kidney, but a breakdown of the material into seasons and areas gave a varying degree of significance. MT in willow ptarmigan may be an important mechanism for detoxifying Cd, and populations exposed to high load may “respond” to the loads by increasing MT synthesis in tissues such as liver and kidney. There is no evidence that willow ptarmigan from areas with high natural Cd loads have reached a limit for MT synthesis in either liver or kidney.     

Patterns of Solidago altissima ramet growth and mortality: the role of below-ground ramet connections

Summary For the rhizomatous perennial, Solidago altissima, I identified clonal fragments in the field, mapped ramet spatial locations, and documented patterns of ramet recruitment, growth, and mortality. Parent ramet size influenced the size and number of daughter ramets produced, and small ramets had lower survivorship and fecundity than large ramets. Similarly, small rhizomes tended to develop into small ramets, and ramets that survived to produce daughter ramets had longer parent-daughter rhizome connections than ramets that did not survive. In addition, most ramets that died during the growing season were connected to (genetically identical) ramets that persisted. There were large size inequalities among rhizomes, ramets, and clonal fragments. Inequalities in the size of ramets increased during the early part of the growing season, then decreased at the end of the season; similar patterns were observed for the growth of clonal fragments. In both instances, the decrease in size inequality could be attributed to the mortality of small individuals (ramets or clonal fragments). I found little evidence that ramet size hierarchies were structured by intraspecific competition. For example, path analyses and randomization tests indicated that size variation among S. altissima ramets was influenced little by the size of their near neighbors (but was influenced by parent size and rhizome size). In addition, within-season variation for the relative size and growth rate of individual ramets led to poor correlations between early and final ramet size; this result suggests that there was no stable hierarchy of dominant and suppressed ramets. I discuss implications of my results for contrasting interpretations of clonal plant population dynamics.     

濒危物种胡杨和灰叶胡杨的克隆生长特征

研究不同生态环境条件下胡杨和灰叶胡杨横走侧根的空间分布及其形态特征、不定芽及克隆分株的数量格局特征及其与土壤水分的关系。研究结果表明:(1)16团3个生境中灰叶胡杨克隆分株所在横走侧根均分布在10—20cm的土层;阿瓦提混交林内灰叶胡杨和胡杨克隆分株所在横走侧根分布在20—40cm的土层;轮台县河道边胡杨克隆分株所在横走侧根分布在10—20cm的土层,林内胡杨克隆分株所在横走侧根分布在20—40cm的土层。不定芽、克隆分株所在部位横走侧根最粗,向两端逐渐变细,且远端根始终比近端根要粗;(2)不定芽具有以前期克隆分株为中心向两端延伸发生分布的空间格局特征,横走侧根上不定芽数与出土克隆分株数呈极显著正相关,未出土克隆分株数与出土克隆分株数呈显著正相关,说明不定芽、未出土克隆分株的数量都直接影响出土克隆分株的数量。(3)不定芽、未出土克隆分株和出土克隆分株出现频率、空间分布范围在同一生境同一根段均表现为不定芽未出土克隆分株出土克隆分株,在不同研究区表现为16团(地下水位1.08m)轮台县(地下水位2.56m)阿瓦提县(地下水位3.34m);相关分析显示土壤水分含量与不定芽和出土克隆分株数量均呈极显著正相关。综合分析认为,地下水位和土壤含水量均对不定芽发生以及不定芽向克隆分株转化有显著的影响。     

Ramet performance in two tussock plants — Do the tussock-level parameters matter?

Studies in plant demography are primarily done at the level of ramets and typically collect ramet-related parameters such as ramet size, type and history. This approach ignores possible effects of factors associated with higher levels, such as genet or tussock. This is particularly important in perennial resprouting herbs with persistent root that consists of many ramets as interaction between ramets, both by competition and by resource sharing, are likely to be intense in these plants. This study investigates effects of tussock-level parameters (age, size and ramet position within tussock and ramet density) on performance of individual ramets in two tussock-forming resprouting herbs (Tanacetum vulgare and Centaurea jacea).The results show that position of a ramet within tussock did not affect ramet growth, but had significant effects on flowering and survival in both species. The direction of the effect differed between the two species; marginal ramets were more successful in T. vulgare, while central ramets were more successful in C. jacea. In addition, tussock age had a significant effect on ramet flowering and survival in T. vulgare. Both these effects are likely to be due to the more competitive life form of T. vulgare, which is restricted to temporary habitats with intense competition. C. jacea is a species of mown or grazed grasslands with lower productivity, which are more stable and where competition is weaker. The effects of the number of neighboring ramets and of the tussock size were significant, but often locality-specific.The results indicate that although many important effects of tussock-related parameters exist, their direction and magnitude differ between species or even populations and are thus not easily predictable. Neglecting these effects, however, is likely to make the demographic models weaker.