Neighbor effects on germination, survival, and growth in two arctic tundra plant communities

In relatively harsh environments such as arctic tundra, abiotic factors have traditionally been considered the primary determinants of community structure, overwhelming any effects of biotic interactions such as competition. Two common low arctic tundra types that differ in soil properties, moist acidic and moist non-acidic tussock tundra (MAT and MNT, respectively), occur in close proximity in northern Alaska. Several plant species occur in both communities with different relative abundance, while others are restricted to one. This study experimentally examined how neighboring vegetation affects germination, survival, and growth of species in these two communities that differ in soil pH, cation availability, and other characteristics. Germination of sown seeds was greater than background levels suggesting seed limitation may restrict recruitment of these clonal, perennial species. Germination of sown seeds was greater at both sites when both mosses and vascular plants had been removed compared to plots with intact vegetation. However, neighbors had almost no effect on survival and growth of adult transplants. Patterns of germination, survival and growth of several species differed depending on the community of origin and the community of destination of the seeds or transplanted adults. For example, transplants of the sedge Eriophorum vaginatum grew better if they were from MAT, and this species germinated better when sown at MNT. Although of relatively short duration (three growing seasons), this study suggests that biotic interactions may affect local species composition by restricting germination and establishment in these two communities, but have less of an effect on adult plants. Not surprisingly, site-specific abiotic conditions also exhibit control over species occurrence and relative abundance. Without disturbance to clear bare ground for recruitment of new individuals, these populations for the most part must rely on clonal growth to persist.     

Growth and tillering patterns within tussocks of Eriophorum vaginatum

Growth patterns were investigated for Eriophorum vaginatum tussocks from disturbed and undisturbed tussock tundra at two sites in Alaska. Total basal area of tussocks decreased with increased cryoturbation but mean basal area per tussock did not. Flowering was observed in tussocks of significantly smaller size on disturbed compared to undisturbed tundra. For tussocks with < 10% cover by shrubs and moss, number of tillers per tussock was linearly related to tussock diameter in most disturbed and undisturbed sites. Exceptions occurred in an area that had been bladed with a bulldozer 7 yr before our survey where tiller number increased as the square of diameter and in an area with much frost activity where tiller number was not emulated with diameter. The ratio of daughter tillers to adult tillers decreased with diameter in disturbed tundra, whereas the trend was less pronounced in undisturbed tundra.
Microsuccession in undisturbed tussock tundra was investigated by sampling tussocks with different amounts of shrub and moss cover for number of daughter tillers per adult tiller, weight per tiller, percent nitrogen, and percent phosphorus. Small tussocks without cover by other species and large, partially covered tussocks were not significantly different by any measures, but tussocks that were almost completely buried had significantly (p < 0.05) lower values of tillering index, weight per tiller, and percent phosphorus.     

Seedling density and seedling survival in Alaskan cotton grass tussock tundra

Previous workers have suggested that reproduction by seedling establishment is less important in the maintenance of tundra vegetation than in temperate areas, but few quantitative data are available. Seedling densities were studied at 11 tundra sites in or near disturbed and undisturbed cotton grass tussock tundra in Alaska, and numerous seedlings were found. Examination of seedling age structures suggests that at least Ledum palustre and probably also Empetrum nigrum do replace themselves by seed in undisturbed tundra. Seedling densities vary greatly depending on the substrate available for establishment.     

DISTURBANCE,LIFE HISTORY STRATEGIES,AND SEED FATES IN ALPINE HERBFIELD COMMUNITIES

Species responses to disturbance are governed primarily by their life history and physiological traits and by the characteristics of the disturbance. Species reproductive traits are especially important in determining the potential of species to establish and to persist following disturbance. Herein, I review available literature on relationships among disturbance, species life histories, and seed fates in tundra environments. Research conducted on these relationships in alpine herbfield vegetation on the Beartooth Plateau, Montana, over the past 9 yr by my colleagues and myself is synthesized. In tundra environments, species reproductive capacities are often similar to those in more temperate environments, but short, cool growing seasons constrain seed production and reduce seedling growth and survival. Highly variable growing season conditions result in large differences in seed production and seedling establishment among years. On disturbed sites, disturbance characteristics determine the seed and seedling environment and influence rates of establishment. In these windy environments, relationships among soil surface characteristics and seed morphological attributes determine both the horizontal and vertical movement of seeds on exposed soils. Once seeds are incorporated into the soil, soil physical and chemical properties determine temperature and nutrient regimes and have the greatest effects on seed germination and seedling growth and survival. Examining the seed fates of herbfield species with varying life histories illustrates that the identities of species that establish following disturbance are largely predictable from their reproductive traits. Disturbance characteristics determine the success of different reproductive strategies and significantly influence community structure.     

Seasonal dynamics and standing crops of biomass and nutrients in a subarctic tundra vegetation

Standing crops of biomass and nutrients were measured in Eriophorum vaginatum tussock tundra and on a north-facing slope, called the camp site, with similar species composition during the summer of 1976 at Eagle Creek, Alaska. These data were then compared to similar data collected at Meade River, Alaska in 1975. Four species are compared: Ledum palustre, Salix pulchra, Betula nana , and Eriophorum vaginatum . The density of aboveground individuals was greater at the tussock site than at the camp site. The total late season above- and belowground standing crop of organic matter and of biomass was greater at the camp site. The nitrogen and calcium contents of new leaves usually increased during the season while phosphorus and potassium contents decreased. Most of the nutrients were in the mosses and lichen compartments rather than in vascular plants.     

种子异型植物异子蓬的生殖分配和结实格局

选择能产生异型果实和种子的一年生盐生植物异子蓬(Suaeda aralocaspica)为研究对象, 研究其生殖分配和结实格局。结果表明, 异子蓬具有较高的生殖分配和规律性的“谨慎型”结实格局。异子蓬的生殖分配高达56%。在资源充裕时, 该植物对扁圆形棕色种子(采取机会主义的萌发策略)的投资多于双凸镜形黑色种子(采取谨慎的萌发策略)的投资。异子蓬的花序类型为二歧聚伞花序, 单个果序含果实(种子)的数目为1-15个, 最多可分为4级。果序内果实的分布具有一定的规律: 第一级1个果实, 第二级2个, 第三级4个, 第四级8个。异子蓬优先将资源供给黑色种子的果实, 利用不同类型种子的发育顺序, 首先保证黑色种子的产出。具有较高的生殖分配和采取“谨慎”的生殖策略是异子蓬与其所处盐漠环境长期适应的结果。     

The response of Arctic vegetation and soils following an unusually severe tundra fire

Fire causes dramatic short-term changes in vegetation and ecosystem function, and may promote rapid vegetation change by creating recruitment opportunities. Climate warming likely will increase the frequency of wildfire in the Arctic, where it is not common now. In 2007, the unusually severe Anaktuvuk River fire burned 1039 km² of tundra on Alaska''s North Slope. Four years later, we harvested plant biomass and soils across a gradient of burn severity, to assess recovery. In burned areas, above-ground net primary productivity of vascular plants equalled that in unburned areas, though total live biomass was less. Graminoid biomass had recovered to unburned levels, but shrubs had not. Virtually all vascular plant biomass had resprouted from surviving underground parts; no non-native species were seen. However, bryophytes were mostly disturbance-adapted species, and non-vascular biomass had recovered less than vascular plant biomass. Soil nitrogen availability did not differ between burned and unburned sites. Graminoids showed allocation changes consistent with nitrogen stress. These patterns are similar to those seen following other, smaller tundra fires. Soil nitrogen limitation and the persistence of resprouters will likely lead to recovery of mixed shrub–sedge tussock tundra, unless permafrost thaws, as climate warms, more extensively than has yet occurred.     

Diurnal patterns of CO2 and H2O exchange of the Arctic sedges Eriophorum angustifolium and E. vaginatum (Cyperaceae)

Eriophorum vaginatum and E. angustifolium are dominant arctic sedges of the well-drained tussock tundra and the permanently flooded wet-sedge tundra, respectively. We determined diurnal courses of gas exchange and water relations of the two species in their natural habitat and compared their responses to changes in light, air temperature, and humidity. Mean photosynthetic response to light was similar between E. angustifolium and E. vaginatum and carbon gain in both species was light limited during most of the growing season. On sunny and dry days, both species closed stomata in response to high leaf-to-air vapor pressure deficits. Even though E. angustifolium was growing in standing water, it exhibited a tighter control of transpirational water loss and had lower hydraulic conductivity in the soil-root-shoot pathway than E. vaginatum. The different response pattern between the two species is discussed in the context of differences in habitat conditions.     

No allocation trade-offs between flowering and sproutingin the lignotuberous,Mediterranean shrub Erica australis

Trade-offs between allocation to sexual or vegetative regeneration capacity are well established as a driving force in the life history patterns of plants in fire-prone environments. However, it is not known whether such trade-offs exist in plants which after aboveground removing disturbances, such as fire, may regenerate by sexual (seeding) or asexual (sprouting) mechanisms. We evaluated whether in the fire-recruiting resprouter Erica australis, which after fire can regenerate by seedling establishment or resprouting, a larger investment in flowers and seeds prior to being disturbed by clipping its aboveground parts would decrease subsequent sprouting, that is, its vegetative regeneration capacity. We analysed the relationships between flower and seed production and the ensuing production and growth of sprouts of six plants from thirteen different sites in central-western Spain. We found no significant relationships between measures of sexual reproductive effort and resprout production and growth 6 months after clipping the aboveground parts of the plants. No evidence of trade-offs between sexual and asexual efforts was found. Furthermore, no significant relationship was found between lignotuber total non-structural carbohydrates and sexual reproductive effort. In addition, 2 years after the disturbance, resprout biomass was positively and significantly correlated with sexual reproductive effort prior to the disturbance. This indicates that growth of resprouts was higher at the sites where plants made a greater reproductive effort. The sites that were more favourable to producing flowers and seeds could also be more favourable to resprouting.     

Competition for tracer 15N in tussock tundra ecosystems

The objectives of this study were to assess the roles of plant species, time, and site on competition for tracer ¹⁵N (without carrier) in tussock tundra ecosystems. Six experimental sites were located in northern Alaska. After one year across the experimental sites, the recovery of ¹⁵N by litter (11.3–16.3%) and mosses (5.4–16.4%) was significantly greater than for aboveground vascular plants (2.6-5.0%). ¹⁵N recoveries by tundra vascular plants (2.6–5.0%) were low when compared to forest trees (9–25%) which suggest that competition for nitrogen is particularly severe in these cold-dominated tundra ecosystems. There were no significant differences among sites in ¹⁵N recoveries by vascular plants, by mosses, or by litter. There was a statistically significant decline in ¹⁵N recovery with time for Vaccinium vitis-idaea and Eriophorum vaginatum between the second and third year. The shallow rooted Vaccinium vitis-ideae was more highly labeled than the deep rooted Eriophorum vaginatum . Nearness to the source of the applied ¹⁵N played a critical role in competition for surface applied nitrogen.     

Effects of waterlogging and increased soil nutrients on growth and reproduction of Polygonum hydropiper in the hydro-fluctuation belt of the Three Gorges Reservoir Region

水淹和土壤养分是影响三峡库区消落带植物生长的主要环境因子。消落带不同高程的植物长期经历不同的淹水强度和土壤养分条件。该研究假设同一物种来自于消落带不同高程的植株可能产生性状分化, 从而对根部淹水和土壤养分变化具有不同的生长和繁殖响应策略。为了验证以上假设, 选取在三峡库区消落带高低高程均广泛分布的物种水蓼(Polygonum hydropiper)为研究对象, 采集自然种群的种子。在温室同质园条件下, 研究了根部水淹和土壤养分提升对高低高程水蓼植株生长和繁殖特性的影响。研究结果表明根部水淹显著或趋于显著降低了水蓼植株功能叶的叶长、叶宽、总分枝数、叶生物量、花生物量和总生物量; 低养分处理显著或趋于显著降低了水蓼植株的总节数、总分枝数、根生物量、花生物量和总生物量, 表明根部水淹和低土壤养分对水蓼的生长和繁殖能力具有抑制作用。同时, 根部水淹和土壤养分的交互作用显著影响植株的根生物量, 表明根部水淹条件下高土壤养分更有利于植株根生物量的积累。高高程植株的根生物量和叶生物量显著或趋于显著高于低高程植株, 而低高程植株的始花时间早于高高程植株, 且繁殖分配也显著高于高高程植株, 表明高低高程水蓼植株对资源的分配策略不同。该研究结果表明水蓼的生长和繁殖特性受根部水淹和土壤养分共同限制, 但对根部水淹条件下高土壤养分生境具有较好的适应性; 同时, 低高程植株可以通过调整其生长和繁殖特性以提高对所处生境胁迫的适应性。     

Phosphate absorption and root respiration of different plant growth forms from northern Alaska

Rate of phosphate absorption measured on field-collected, excised roots of eight species from a tundra site and a taiga forest was more strongly correlated with growth form traits than with type of community. In the taiga forest phosphate absorption rate ranged tenfold among species and was higher in graminoids than in shrubs. In both sites deciduous shrubs tended to have higher rates than evergreens. Phosphate absorption rate was not strongly reduced at low temperature in any species examined and there was a temperature optimum of about 20°C in Eriophorum vaginatum . Phosphate absorption rates were higher in lateral than primary roots, although both root types had substantial respiration rates. At one tundra site, calculations suggest that roots of E. vaginatum require 30–40 d to absorb a quantity of phosphorus equivalent to that required to synthesize the root and another 5–6 d to provide the net phosphorus requirements for the remainder of the plant.     

根部水淹和土壤养分提升对三峡库区消落带水蓼生长和繁殖特性的影响

水淹和土壤养分是影响三峡库区消落带植物生长的主要环境因子。消落带不同高程的植物长期经历不同的淹水强度和土壤养分条件。该研究假设同一物种来自于消落带不同高程的植株可能产生性状分化, 从而对根部淹水和土壤养分变化具有不同的生长和繁殖响应策略。为了验证以上假设, 选取在三峡库区消落带高低高程均广泛分布的物种水蓼(Polygonum hydropiper)为研究对象, 采集自然种群的种子。在温室同质园条件下, 研究了根部水淹和土壤养分提升对高低高程水蓼植株生长和繁殖特性的影响。研究结果表明根部水淹显著或趋于显著降低了水蓼植株功能叶的叶长、叶宽、总分枝数、叶生物量、花生物量和总生物量; 低养分处理显著或趋于显著降低了水蓼植株的总节数、总分枝数、根生物量、花生物量和总生物量, 表明根部水淹和低土壤养分对水蓼的生长和繁殖能力具有抑制作用。同时, 根部水淹和土壤养分的交互作用显著影响植株的根生物量, 表明根部水淹条件下高土壤养分更有利于植株根生物量的积累。高高程植株的根生物量和叶生物量显著或趋于显著高于低高程植株, 而低高程植株的始花时间早于高高程植株, 且繁殖分配也显著高于高高程植株, 表明高低高程水蓼植株对资源的分配策略不同。该研究结果表明水蓼的生长和繁殖特性受根部水淹和土壤养分共同限制, 但对根部水淹条件下高土壤养分生境具有较好的适应性; 同时, 低高程植株可以通过调整其生长和繁殖特性以提高对所处生境胁迫的适应性。     

Biomass allocation patterns in terrestrial, epiphytic and aquatic species of Utricularia (Lentibulariaceae)

Utricularia forms the largest genus of carnivorous plants and is characterized by the possession of typical traps (“bladders”). Total biomass allocation was examined in three aquatic, six terrestrial and one epiphytic species of Utricularia from natural habitats in West Africa and from the Botanical Gardens, Bonn. Total biomass of aquatic species was considerably higher than that of terrestrial or epiphytic species. Epiphytic Utricularia accumulate about 35% of their biomass in green leaves, in contrast to 65% of nearly chlorophylless reproductive structures and traps. Aquatic species allocated more than 85% of their total biomass to stolons, leaves and traps, but only 10–13% to reproductive structures. This is in stark contrast to the allocation patterns of terrestrial bladderworts. These species allocate nearly 90% of their total biomass in reproductive structures, and only about 10% to stolons, leaves and traps. This reduction of photosynthetically active plant tissue strongly suggests that as a consequence of the alternative resource of chemical energy, the carnivorous habit might have partly replaced autotrophy in certain terrestrial Utricularia species, especially in some smaller ones.     

Sexual differences in biomass and nutrient allocation of first-year Silene dioica plants

Reproductive and somatic biomass, nitrogen (N), and phosphorus (P) pools were compared between females and males in 1st-year plants of Silene dioica. We estimated irretrievable resources allocated to seeds, pollen, flowers, and unrecovered summer leaf investment by collecting plant parts at abscission throughout the season. At the end of the season, we determined resources lost through senescent stems and autumn leaf turnover and resources stored in perennial roots and overwintering buds. Sexual differences in allocation patterns depended on the resource used for comparison, and whether absolute or proportional resource pools were assessed. Total resource pools in terms of biomass and N were similar for females and males. However, male plants acquired relatively more P. The proportional reproductive investment, i.e., reproductive effort, was similar for males and females in terms of biomass and N. In terms of P, male reproductive effort was higher. There was no difference between sexes in the proportional and relative biomass allocated to perennial roots and overwintering buds. However, in terms of absolute and relative N allocation to below-ground parts, females had larger reserves than males. Females, moreover, had a larger proportion of their P in below-ground parts. However, as male total P pools were larger, absolute P reserves did not differ between sexes. The high reproductive effort and N depletion of below-ground parts in males resulted largely from higher flower production compared to females. In females, seeds were the major component of reproductive effort. These results show that if biomass and nutrient allocation are assessed in parallel for dioecious plants, we obtain a more complete view of their sexual differences. Received: 07 May 1998 / Accepted: 30 October 1998     

Inherent allocation patterns and potential growth rates of herbaceous climbing plants

We tested the hypothesis that herbaceous climbing plants, unlike non-climbing herbs, maximize height growth and leaf area, with minimal expenditure in support structures. The enhanced investment in leaf area was expected to result in high relative growth rates in terms of biomass increment. Four leguminous herbaceous climbers from nutrient-poor sites and four non-leguminous herbaceous climbers from nutrient-rich sites, were compared with non-climbing, self-supporting leguminous and non-leguminous herbaceous species from similar habitats. Plants were grown in hydroponic cultures in controlled environment chambers. All climbers had inherently taller shoots than self-supporting plants when compared at an equal amount of total plant dry weight, due to longer stems per unit of support biomass. In contrast to the hypothesis, the relative growth rates of all climbers were relatively low compared to the range found for self-supporting species. The biomass allocation patterns of the non-leguminous climbers were similar to those of the self-supporting species. Leguminous climbers allocated more biomass to support tissue and less biomass to leaves than non-climbers. As a result, height growth was even more emphasized in leguminous climbers than in non-leguminous climbers. Climbing legumes had high rates of net carbon gain, which partly compensated the lower relative leaf weight. We conclude that leguminous herbaceous climbers maximize height growth by a large investment in support biomass, enabling them to keep a large proportion of their leaves in the better illuminated environment at the top of the vegetation canopy.     

Changes in individual plant traits and biomass allocation in alpine meadow with elevation variation on the Qinghai-Tibetan Plateau

Plant traits and individual plant biomass allocation of 57 perennial herbaceous species, belonging to three common functional groups (forbs, grasses and sedges) at subalpine (3700 m ASL), alpine (4300 m ASL) and subnival (⩾5000 m ASL) sites were examined to test the hypothesis that at high altitudes, plants reduce the proportion of aboveground parts and allocate more biomass to belowground parts, especially storage organs, as altitude increases, so as to geminate and resist environmental stress. However, results indicate that some divergence in biomass allocation exists among organs. With increasing altitude, the mean fractions of total biomass allocated to aboveground parts decreased. The mean fractions of total biomass allocation to storage organs at the subalpine site (7%±2% S.E.) were distinct from those at the alpine (23%±6%) and subnival (21%±6%) sites, while the proportions of green leaves at all altitudes remained almost constant. At 4300 m and 5000 m, the mean fractions of flower stems decreased by 45% and 41%, respectively, while fine roots increased by 86% and 102%, respectively. Specific leaf areas and leaf areas of forbs and grasses deceased with rising elevation, while sedges showed opposite trends. For all three functional groups, leaf area ratio and leaf area root mass ratio decreased, while fine root biomass increased at higher altitudes. Biomass allocation patterns of alpine plants were characterized by a reduction in aboveground reproductive organs and enlargement of fine roots, while the proportion of leaves remained stable. It was beneficial for high altitude plants to compensate carbon gain and nutrient uptake under low temperature and limited nutrients by stabilizing biomass investment to photosynthetic structures and increasing the absorption surface area of fine roots. In contrast to forbs and grasses that had high mycorrhizal infection, sedges had higher single leaf area and more root fraction, especially fine roots.     

Environmental control of reproductive phenology and the effect of pollen supplementation on resource allocation in the cleistogamous weed, Ruellia nudiflora (Acanthaceae)

• Background and Aims Mixed reproductive strategies may have evolved as a response of plants to cope with environmental variation. One example of a mixed reproductive strategy is dimorphic cleistogamy, where a single plant produces closed, obligately self-pollinated (CL) flowers and open, potentially outcrossed (CH) flowers. Frequently, optimal environmental conditions favour production of more costly CH structures whilst economical and reliable CL structures are produced under less favourable conditions. In this study we explore (1) the effect of light and water on the reproductive phenology and (2) the effect of pollen supplementation on resource allocation to seeds in the cleistogamous weed Ruellia nudiflora.• Methods Split-plot field experiments were carried out to assess the effect of shade (two levels: ambient light vs. a reduction of 50 %) and watering (two levels: non-watered vs. watered) on the onset, end and duration of the production of three reproductive structures: CH flowers, CH fruit and CL fruit. We also looked at the effect of these environmental factors on biomass allocation to seeds (seed weight) from obligately self-pollinated flowers (CL), open-pollinated CH flowers and pollen-supplemented CH flowers.• Key Results CH structures were produced for a briefer period and ended earlier under shaded conditions. These conditions also resulted in an earlier production of CL fruit. Shaded conditions also produced greater biomass allocation to CH seeds receiving extra pollen.• Conclusions Sub-optimal (shaded) conditions resulted in a briefer production period of CH structures whilst these same conditions resulted in an earlier production of CL structures. However, under sub-optimal conditions, plants also allocated more resources to seeds sired from CH flowers receiving large pollen loads. Earlier production of reproductive structures and relatively larger seed might improve subsequent success of CL and pollen-supplemented CH seeds, respectively.     

新疆郁金香营养生长、个体大小和开花次序对繁殖分配的影响

植物有性繁殖与资源分配的关系研究对于揭示植物生活史特征及繁育系统进化具有重要意义。新疆郁金香(Tulipa sinkiangensis)是新疆天山北坡荒漠带特有的一种多年生早春短命植物。在自然生境中,该物种仅以有性繁殖产生后代,每株能产生1-8朵花,且不同植株上的花数及果实数以及花序不同位置上的花与果实大小明显不同。本文通过对新疆郁金香有性繁殖与营养生长及植株大小的关系以及花序中不同位置花及果实间的资源分配研究,旨在揭示营养生长、个体大小及开花次序对其繁殖分配的影响。结果表明:在开花和果实成熟阶段,新疆郁金香植株分配给营养器官(鳞茎和地上营养器官)与繁殖器官的资源间均存在极显著的负相关关系(P<0.01),说明其植株的营养生长与生殖生长间存在权衡关系。多花是新疆郁金香的一个稳定性状,其植株上花数目、花生物量、果实生物量和种子数量与植株生物量间均呈极显著的正相关关系(P<0.01),说明新疆郁金香植株的繁殖分配存在大小依赖性。在具2-5朵花的新疆郁金香植株中,花序内各花的生物量、花粉数和胚珠数、结实率、果实生物量、结籽数、结籽率及种子百粒重按其开花顺序依次递减,说明花序内各花和果实的资源分配符合资源竞争假说。植株通过减少晚发育的花或果实获得的资源来保障早发育的花或果实获得较多的资源,从而达到繁殖成功。