Above-ground competition does not alter biomass allocated to roots in Abutilon theophrasti

We tested whether plants allocate proportionately less biomass to roots in response to above-ground competition as predicted by optimal partitioning theory. Two population densities of Abutilon theophrasti were achieved by planting one individual per pot and varying spacing among pots so that plants in the two densities experienced the same soil volume but different degrees of canopy overlap. Density did not affect root:shoot ratio, the partitioning of biomass between fine roots and storage roots, fine root length, or root specific length. Plants growing in high density exhibited typical above-ground responses to neighbours, having higher ratios of stem to leaf biomass and greater leaf specific area than those growing in low density. Total root biomass and shoot biomass were highly correlated. However, storage root biomass was more strongly correlated with shoot biomass than was fine-root biomass. Fine-root length was correlated with above-ground biomass only for the small subcanopy plants in crowded populations. Because leaf surface area increased with biomass, the ratio between absorptive root surface area and transpirational leaf surface area declined with plant size, a relationship that could make larger plants more susceptible to drought. We conclude that A. theophrasti does not reallocate biomass from roots to shoots in response to above-ground competition even though much root biomass is apparently involved in storage and not in resource acquisition.     

Influence of intraspecific interaction and substrate type on initial growth and establishment of Hydrilla verticillata

Both substrate type and plant–plant interaction can greatly influence the growth and establishment of plants. In order to assist the re-vegetation of submerged macrophytes, the growth of Hydrilla verticillata with increasing equi-distance neighboring plant density on two substrate types (sediment and sand, representing high- and low-nutrient level, respectively) was assessed in monoculture stands. The results showed that substrate type greatly changed the biomass allocation patterns of the target plants, with a smaller root mass ratio on sediment compared to sand (0.70 vs. 3.11%). However, interaction between substrate type and neighboring density was observed. At low density, growth on sediment greatly increased plant height (43.90 vs. 22.10 cm), leaf biomass (216.63 vs. 68.41 mg), and total biomass (298.39 vs. 121.77 mg) when compared to growth on sand. However, at high density, no significant effect of the substrate type was found in those parameters. On sediment, high neighboring density greatly decreased the height, root number, total root length, root mass, and total biomass, implying large intraspecific plant–plant competition. However, such competition can be greatly reduced in infertile environments. Therefore, when the plants were grown on sand, neighboring density showed little effect on the height (22.10–26.53 cm), total root length (21.34–40.50 cm), and root biomass (3.14–6.27 mg). Total biomass and root number significantly increased by 50% and 115%, respectively, at high density compared to low density on sand, suggesting that facilitation rather than competition was occurring. Therefore, plant–plant interaction can vary from competition in fertile environments to facilitation in infertile environments. In summary, neighboring density should be manipulated according to the environmental nutrient level, so as to reduce intraspecific competition or increase intraspecific facilitation, and finally enhance the initial growth and establishment of H. verticillata in re-vegetation activities.     

Increased root oxygen uptake in pea plants responding to non-self neighbors

Recent studies have demonstrated that plants alter root growth and decrease competition with roots of the same individual (self); however, the physiological traits accompanying this response are still widely unknown. In this study, we investigated the effect of root identity on gas exchange in the model species pea (Pisum sativum L.). Split-root plants were planted so that each pot contained either two roots of the same plant (self) or of two different plants (non-self), and the responses of biomass, photosynthesis, and respiration were measured. The photosynthetic rate was not affected by the identity of the root neighbor. We found a reduction of leaf dark respiration by half, accompanied by an increase in nocturnal root respiration by 29 % in plants neighboring with non-self. The activity of the alternative oxidase (AOX) pathway increased when plants responded to non-self neighbors. The increased activity of AOX in plants responding to non-self indicates carbon imbalances in roots, possibly as a consequence of increased root exudation and communication between individuals. If such an effect occurs more widely, it may change the assumptions made for the quantity of respiration as used in carbon budget models.     

杉木成熟林细根形态与功能特征的海拔梯度变异特点

为探究植物对环境变化的适应策略,在安徽省金寨县天马国家自然保护区,以不同海拔高度(750、850、1000、1150 m)杉木(Cunninghamia lanceolata)成熟林为对象,采用土钻法获取土壤细根样品,分别测定了不同海拔不同土层(0—10 cm、10—20 cm、20—30 cm)土壤细根生物量、形态特征参数和碳氮含量。结果表明:(1)随海拔梯度增加,0—30 cm土层细根生物量、根长密度、比根长、表面积密度、体积密度均呈先减少后增加趋势,在海拔750 m生物量最大,其余指标在海拔1150 m最大;随土层深度增加,同一海拔细根生物量、根长密度、表面积密度、体积密度均呈减少趋势。(2)随海拔梯度增加,0—30 cm土层细根C和N含量呈先增加后减少趋势,C/N比呈先减少后增加再减少趋势;随土层深度增加,同一海拔细根C含量呈先减少后增加趋势,N含量呈降低趋势,C/N比呈上升趋势。(3)细根N含量与生物量、根长密度和体积密度显著正相关,C/N比与生物量、根长密度、表面积密度和体积密度极显著负相关。(4)土壤水分对细根生物量及其形态指标影响显著。     

Metal complexes increase uptake of Zn and Cu by plants: implications for uptake and deficiency studies in chelator-buffered solutions

The allocation of resources among roots and shoots represents the largest flux of resources within a plant and therefore should have been selected to maximize benefits to plants. Yet, it is unclear why some species like temperate grasses have such high root length density (RLD). Either the slow rate of diffusion of inorganic N in soils or interplant competition could explain the high RLD of temperate grasses. Using a fine-scale model of nutrient dynamics in the soil and plant growth, a cost–benefit approach was used to assess optimal allocation rates for plants that accounted for value of both carbon and nitrogen. In the absence of interplant competition, resource benefits are maximized with very little root length except in extremely dry soils for ammonium. In the presence of a competitor, optimal allocation of N to roots is much greater and increases as ability of competitors to produce root length increase. Competition for inorganic nitrogen generates a classic aspect of the tragedy of the commons, the “race for fish”, where plants must allocate more resources to acquisition of the limiting resource than is optimal for plants in the absence of competition. As such, nutrient competition needs to be directly addressed when understanding plant- and ecosystem-level resource fluxes as well as the evolution of root systems.     

Aggregative Root Placement: A Feature During Interspecific Competition in Inland Sand-Dune Habitats

Segregation of roots is frequently observed in competing root systems. However, recently, intensified root growth in response to a neighbouring plant has been described in pot experiments [Gersani M, Brown J S, O'Brien E E, Maina G M and Abramsky Z 2001. J. Ecol. 89, 660–669]. This paper examines whether intense root growth towards a neighbour (aggregation) plays a role in competitive interactions between plant species from open nutrient-poor mid-European sand ecosystems. In a controlled field-competition experiment, root distribution patterns of intra- and interspecific pairs as well as single control plants of Corynephorus canescens, Festuca psammophila, Hieracium pilosella, Hypochoeris radicata and Conyza canadensis were investigated after one growing season. Under intraspecific competition plants tended to segregate their root systems, while under interspecific competition most species tended to aggregate roots towards their neighbours even at the expense of root development at the opposite competition-free side of the target. Preference of a root aggregation strategy over the occupation of competition-free soil in interspecific competition emphasizes the importance of contesting between individuals in relation to mere resource acquisition. It is suggested that in the presence of a competitor the plants might use root aggregation as a defensive reaction to maintain a strong competitive response and exclusive access to the resources of already occupied soil volumes.     

Intra-plant versus Inter-plant Root Competition in Beans: avoidance,resource matching or tragedy of the commons

Root competition inhibits root proliferation. All else equal, a plant should invest roots in a nutrient patch devoid of roots rather than one already occupied by roots. Less clear is how a plant should respond to intra-plant versus inter-plant root competition. We consider three responses for how a plant may select habitats based on intra-versus inter-plant root competition: inter-plant avoidance, resource matching, or intra-plant avoidance. The first assumes that plants prefer to have their own space and preferentially proliferate roots away from neighboring plants. The second response, based on the ideal free distribution, assumes that plants invest so as to equalize average returns from roots, regardless of the identity of the neighboring roots. The third, based on game theory, assumes that the plant proliferates roots so as to maximize whole-plant fitness, in which case it is better to proliferate plants among a neighbor's roots than to continue proliferating amongst one's own roots. To test among these models we grew beans (Phaseolus varigaris, var. Kenya) in a greenhouse under two planting scenarios. Both scenario were tested under 0.5 and 0.1 strength of nutrient solution. Under scenario A (fence-sitters), two split-root plants each shared two patches by virtue of having roots in each. Under scenario B (owners) two plants each had their own patch. The results supported the game theory model of intra-plant avoidance (whole plant habitat selection). Fence-sitters produced 150% more root mass per individual than owners. Owners produced 90% more yield (dry mass of pods) than fence-sitters. Furthermore, owners had significantly higher shoot-root ratios than fence-sitters. These effects did not vary with high or low nutrient levels. The over-proliferation of roots under inter-plant competition (fence-sitters) was manifest by the tenth day after planting. In short, the fence-sitters engaged in a tragedy of the commons in which they competed with each other through root proliferation. At the ESS, the fitness maximizing strategy of the individual is to sacrifice collective yield in a quest to `steal' nutrients from its neighbor. The research has three implications. First, plants may be able to assess and respond to local opportunities in a manner that maximizes the good of the whole plant. Second, nutrient foraging as a game may provide a fresh perceptive for viewing root competition either intra-specifically or inter-specifically. Third, it may be possible to increase the yield of certain crop species by breeding more `docile' cultivars that do not overproduce roots in response to inter-plant competition.     

喜旱莲子草和接骨草竞争对模拟增温的响应北大核心CSCD

研究外来入侵植物与本地植物种竞争对气候变暖的响应,对于预测未来气候变化背景下入侵植物的入侵趋势、理解其入侵机制以及筛选生态替代种具有重要的意义。以入侵我国的外来植物喜旱莲子草(Alternanthera philoxeroides)和本地植物种接骨草(Sambucus chinensis)为材料,通过两种植物单栽、纯栽和混栽,采用红外辐射加热器模拟增温,研究了两种植物竞争对模拟增温的响应。结果表明:(1)在模拟增温期间(2013年5–12月),增温组空气平均温度比不增温组提高了0.47℃,相对湿度降低了1.87%;(2)混栽的喜旱莲子草除根冠比与单栽无显著差异外,其余各生物量和根系形态指标均显著低于单栽喜旱莲子草;无竞争、种间竞争和种内竞争三种竞争间,接骨草除根冠比、细根与总根生物量比、比根长和比根表面积无显著差异外,其余指标均呈现无竞争>种间竞争>种内竞争的趋势;(3)无竞争、种间竞争和种内竞争三种条件下,喜旱莲子草各指标在增温和不增温处理间差异均不显著,而接骨草总生物量和根生物量在无竞争和种间竞争条件下增温处理均显著低于不增温处理,在种内竞争条件下则相反;(4)增温使接骨草的相对拥挤系数降低,接骨草对温度升高反应敏感,而喜旱莲子草则表现出一定的适应性。由此推测,在中度遮阴陆生生境中,接骨草有望成为喜旱莲子草生物替代控制的材料。     

Mechanisms for the increase in phosphorus uptake of waterlogged plants: soil phosphorus availability, root morphology and uptake kinetics

Waterlogging frequently reduces plant biomass allocation to roots. This response may result in a variety of alterations in mineral nutrition, which range from a proportional lowering of whole-plant nutrient concentration as a result of unchanged uptake per unit of root biomass, to a maintenance of nutrient concentration by means of an increase in uptake per unit of root biomass. The first objective of this paper was to test these two alternative hypothetical responses. In a pot experiment, we evaluated how plant P concentration of Paspalum dilatatum, (a waterlogging-tolerant grass from the Flooding Pampa, Argentina) was affected by waterlogging and P supply and how this related to changes in root-shoot ratio. Under both soil P levels waterlogging reduced root-shoot ratios, but did not reduce P concentration. Thus, uptake of P per unit of root biomass increased under waterlogging. Our second objective was to test three non-exclusive hypotheses about potential mechanisms for this increase in P uptake. We hypothesized that the greater P uptake per unit of root biomass was a consequence of: (1) an increase in soil P availability induced by waterlogging; (2) a change in root morphology, and/or (3) an increase in the intrinsic uptake capacity of each unit of root biomass. To test these hypotheses we evaluated (1) changes in P availability induced by waterlogging; (2) specific root length of waterlogged and control plants, and (3) P uptake kinetics in excised roots from waterlogged and control plants. The results supported the three hypotheses. Soil P avail-ability was higher during waterlogging periods, roots of waterlogged plants showed a morphology more favorable to nutrient uptake (finer roots) and these roots showed a higher physiological capacity to absorb P. The results suggest that both soil and plant mechanisms contributed to compensate, in terms of P nutrition, for the reduction in allocation to root growth. The rapid transformation of the P uptake system is likely an advantage for plants inhabiting frequently flooded environments with low P fertility, like the Flooding Pampa. This advantage would be one of the reasons for the increased relative abundance of P. dilatatum in the community after waterlogging periods. Received: 15 February 1997 / Accepted: 20 May 1997     

Effects of density on above and below ground biomass of the native alpine grass Poa fawcettiae and the environmental weed Achillea millefolium

Competition between Poa fawcettiae Vickery, the dominant native snowgrass, and the invasive herbaceous Achillea millefolium L., was examined in three glasshouse experiments. The first experiment investigated the potential for intra-specific competition in plants by growing them in pots with low and high density. The second experiment examined the potential for inter-specific competition at low, medium and high density. In the third experiment plants in pots where either roots or shoots of the species could not compete were compared to those where root and shoot competition was possible.
Achillea millefolium plants produced more than four times the biomass of P. fawcettiae plants. As a result the two species responded differently. In the A. millefolium monocultures both root and shoot biomass per plant declined at high density. By contrast, P. fawcettiae biomass was not affected. In mixed species pots, P. fawcettiae had no effect on the biomass of A . millefolium plants, while P. fawcettiae shoot and root biomass per plant decreased when grown with A. millefolium at all densities tested. Root competition from A. millefolium appears to be the main cause of the decrease in biomass of P. fawcettiae . The results imply that A. millefolium may have a competitive advantage over P. fawcettiae in the Australian Alps.     

不同生长阶段苘麻生物量分配对种群密度和土壤水分的响应

通过研究不同生长阶段下植物生物量分配对土壤水分和种群密度的响应,揭示植物同时应对生物与非生物环境因子的策略。本研究在田间条件下对1年生草本植物苘麻（Abutilon theophrasti）进行加水和对照2种水分处理,每种处理下进行低、中、高3种种植密度处理,分别在生长20、50 d时测量植物根、茎、叶片、叶柄和繁殖（花和果实）生物量,探讨在不同生长阶段苘麻生物量分配如何响应于密度和水分。结果表明：植物生长20 d时,在加水处理中,与低密度相比,中密度提高了根生物量比率19.4%和根冠比21.5%,降低了叶生物量比率34.4%;未加水处理（对照）中生物量分配对密度的响应不显著;50 d时,对照处理下,高密度相对于低密度降低了总生物量63.5%,2种水分处理下高密度都降低了根冠比和根生物量比率,提高了茎生物量比率,不影响总生物量和其他器官生物量分配。结果说明施加水分前期更容易促进根生物量分配对密度的积极响应（增大）,后期减缓高密度对总生物量的不利影响（降低）。生物量分配对密度的响应取决于种内相互作用的强度,早期适中水平的相互作用更容易产生地下促进作用,促进根部的积极响应。中密度下适中的种...     

The effect of neighbors on the root system of the desert annualEremobium aegyptiacum

Intra-specific competition with a primary focus on root competition between plants living in an area with low resource levels, was studied using the natural monotypic population of a desert annual plantEremobium aegyptiacum (Cruciferae). We tested the effect of neighboring distance on shoot and root biomass, and such root parameters as root length, diameter of root neck, number of first order lateral roots and number of lateral roots per unit of main root length. Our results indicate a strong negative relationship between neighboring plant density and such plant parameters as shoot and root biomass, and root neck diameter. The number of first order lateral roots and the number of lateral roots per unit of main root length were negatively related to the distance between neighboring plants. Contrary to predictions, there was no influence of competition on node distribution: neither root overlap nor root avoidance was observed in pairs of adjacent plants.     

Effect of root competition and shading on resprouting dynamics of Erica multiflora L.

Abstract. Resprouting from underground structures is one of the main regeneration strategies of Mediterranean shrubs after aerial biomass disturbance such as fire or clear-cutting to reduce fire risk. In order to study the effect of root competition and shading (simulated shoot competition) on Erica multiflora, growth, morphology, flowering performance and sprout size variability during resprouting, a factorial field experiment was conducted in which neighbours around target plants were eliminated and plants were shaded with mesh for two years. Root competition reduced sprout recruitment and sprout density (number of sprouts per unit stump area) more strongly than did shading. The negative effect of root competition on sprout biomass was constant with time, while the reduction due to shading increased with time. There was an interaction between root competition and shading on the biomass of sprouts 22 months after treatment: genets without root competition and shading were four times larger than in any other treatment. Both shading and root competition also decreased percentage branching but did not modify maximum sprout height. Only shading decreased the leaf/shoot biomass ratio and the percentage of flowering genets. One year after resprouting, root competition counteracted the effect of shading on inducing sprout biomass variability within the genet because it decreased sprout density. 22 months after treatment, sprout biomass variability was not affected by any main effect. The results suggest that competition among sprouts within the genet is asymmetric. However, shading by genet neighbours may not always increase sprout biomass variability if root competition is also severe.     

根系隔离条件下的谷子亲缘识别

在根系隔离情况下, 通过研究邻株身份(亲缘株、非亲缘株、陌生株)及其与种植密度(高、低)和土壤养分水平(高、低)交互作用对谷子(Setaria italica)形态学特征和生物量分配的影响, 探索谷子地上部分是否能够识别亲缘邻株, 以及谷子的这种亲缘识别能力对环境因子如何响应。结果显示: 1)亲缘组谷子叶生物量分配显著降低, 茎秆显著增粗, 暗示着亲缘组谷子植株间减少竞争, 并增强对当地多风气候的适应。而非亲缘组谷子叶生物量分配显著增加, 表明非亲缘组谷子植株间竞争较强。2)与非亲缘组相比, 陌生组谷子种子生物量分配显著增加, 株高显著减少, 表明陌生组谷子植株通过不对称竞争(与邻株糜(Panicum miliaceum)植株相比, 株高显著增加), 进一步限制邻株(糜)生长, 从而增强竞争能力, 同时, 将更多的生物量投资于繁殖, 增加适合度。3)在高密度种植条件下, 谷子茎生物量和叶生物量分配在各邻株身份处理间无显著差异, 而在低密度种植条件下, 与非亲缘组相比, 亲缘组谷子茎生物量显著增加, 叶生物量分配显著减小; 随着种植间距的增大(种植密度减小), 亲缘组谷子叶生物量分配显著减少, 而非亲缘组和陌生组叶生物量分配在高、低种植密度条件下无显著差异。4)在低土壤养分条件下, 亲缘组和非亲缘组谷子叶生物量分配无显著差异, 前者穗长显著小于后者, 而在高土壤养分条件下, 亲缘组谷子叶生物量分配显著小于非亲缘组, 前者穗长显著大于后者。结果表明, 在根系隔离的情况下,谷子能够识别亲缘邻株, 且谷子地上部分竞争信号在亲缘识别过程中扮演重要角色。较低种植密度和较高土壤养分水平有利于谷子亲缘识别能力的表达。     

Kin recognition in Setaria italica under the condition of root segregation

Aims Kin recognition may play an important part in the performance and productivity of crop plants. However, so far, little is known about whether crop plants can recognize their kin neighbors. The aim of this study was to explore kin recognition in Setaria italica, and its responses to changes in environmental and biological conditions.Methods A field experiment was conducted in the suburb of Shanghai. Setaria italica grew with different neighbors (kin, non-kin and strangers), under the condition of root segregation and different plant densities (high and low) and soil nutrient levels (high and low), respectively. We investigated how neighbor identity and its interactions with plant density and soil nutrient level affected the morphology and biomass allocation of S. italica.Important findings Under the condition of root segregation, 1) Leaf biomass allocation and stem diameter of plants in the kin groups significantly decreased and increased, respectively, suggesting that plants of S. italica in the kin groups reduced inter-individual competition, and adapted to the local windy climate. 2) Compared with the non-kin groups, plants in the stranger groups significantly increased the biomass allocation to seeds, while plant height decreased significantly, suggesting that the plants of S. italica in the stranger groups may reduce the growth of their neighbors due to asymmetric competition (S. italica significantly increased height compared with the neighboring plants, Panicum miliaceum). Therefore, the S. italic plants in the stranger groups allocated more biomass to reproduction and increased fitness than those in non-kin groups. 3) Under the condition of high plant density, no significant differences were found in stem biomass and leaf biomass allocation of plants among different neighbor identity treatments. While under the condition of low plant density, compared with the non-kin groups, biomass allocation to stem and leaf in the kin groups significantly increased and decreased, respectively. As the plant density decreased, plants in the kin groups decreased leaf biomass allocation significantly, while plants in the non-kin and stranger groups did not show such a response. 4) Under the condition of low soil nutrient level, no significant difference was found in leaf biomass allocation between the kin and non-kin groups, while the ear length of plants in the kin groups decreased significantly. Under the condition of high soil nutrient level, the biomass allocated to leaves in the kin groups decreased significantly, while ear length increased significantly compared with the non-kin groups. Therefore, under the condition of root segregation, plants of S. italica showed the ability to recognize their kin neighbors, and the aboveground competitive cues may play important roles in the course of kin recognition in S. italica. Lower plant density and higher soil nutrient level may facilitate the ability of kin recognition in S. italica.     

Response of cowpea to variations in planting density and nutrient level

A greenhouse study was carried out using cowpea (Vigna unguiculata (L.) Walp.) grown in Perlite^® and inoculated with Nitragin^® to investigate the concentration of plant nutrients and planting density required for optimum biomass production. Five concentrations (full, 0.5, 0.2, 0.1 and 0.05 strength) of Bisseling's nutrient solution and five planting densities (one to five plants per pot) were tested in a factorial randomized Graeco-Latin square design. Growth was determined as fresh and dry weights of leaves, stems, petioles, roots, flowers and pods, and whole plant.Optimum biomass production was found at 0.5 strength nutrient solution and a density of one plant per pot. Plants were more sensitive to higher planting density than to alterations of nutrient level. Over a twenty-fold range of nutrient supply, whole plant biomass yield varied at most by 44%, whereas increasing planting density from one to five plants per pot decreased biomass production by as much as 77%. There is a decrease in the shoot/root ratio as nutrient level decreases. The data suggests a potential for higher seed production at the higher densities and lowest nutrient levels, but this data was inconclusive.     

Density-dependent responses of Picea purpurea seedlings for plant growth and resource allocation under elevated temperature

This study was conducted to determine whether plants in the presence or absence of competition differ in their responses to warming, and whether density modifies the effect of warming. Picea purourea seedlings were grown under ambient and warming (ambient +2.2 °C) conditions in climate control chambers with two different planting densities. After 4 years, seedlings were harvested and measured for height, stem diameter, leaf area, structural biomass, carbon, nitrogen, chlorophyll and carbohydrate levels of needles, branches, stem and roots. At low density, warming increased height, stem diameter, total leaf area biomass production and carbohydrate concentration per seedling, while it decreased C/N ratio for all plant parts, but did not affect chlorophyll content. By contrast, at high density, although warming increased biomass and total leaf area, it did not affect plant height and stem diameter. At the same time, it had different effects on chlorophyll content, C/N ratio and carbohydrate levels among plant parts. On the other hand, high density limited plant growth and altered resource allocation pattern. Our study demonstrates that planting densities decreased the temperature-induced growth enhancement of P. purpurea seedlings and the effects of warming on resource allocation not only showed density-dependence, but also vary with tissue age classes and root diameter; the responses of plants to elevated temperature, acquired from plants growing as individuals, may not be applicable to plants grown under intraspecific competition as typically found in the field.     

FITNESS CONSEQUENCES OF MULTIPLE PATERNITY IN WILD RADISH,RAPHANUS SATIVUS

In natural populations, wild radish plants typically mate with 6–8 pollen donors, and seeds of individual fruits are usually sired by 1–4 fathers. Since radish fruits are indehiscent and gravity-dispersed, progeny are most likely to compete with a mixture of full and half siblings. The fitness consequences of single and multiple paternity were investigated in a greenhouse experiment. Seeds of every possible cross in a 5 times 5 reciprocal diallel mating design were assigned to one of three competition regimes (four full siblings, four maternal half siblings, or four unrelated individuals per pot) or were grown as singletons. After 14 weeks, the aboveground biomass of all plants was harvested and oven-dried. The dry weight of singletons was more than three times that of progeny grown in competition, indicating that intraspecific competition had occurred. Full- and half-sib progenies did not differ in mean dry weight. Thus, there was no evidence that multiple paternity enhances this aspect of maternal fitness. However, the competition regime dramatically affected the coefficient of variation in dry weight of progeny within a pot. Weight hierarchies were much more pronounced in pots of half sibs and unrelated neighbors than in pots of full sibs. Also variance in dry weight attributable to sire was greatest in the half-sib and “unrelated neighbors” competition regimes. These results suggest that weight hierarchies reinforce genetic differences among the offspring.     

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.     

Rooting strategy of naturally regenerated beech in Silver birch and Scots pine woodlands

This study investigated the belowground development and strategy of late-successional European beech (Fagus sylvatica L.) in ageing natural Scots pine (Pinus sylvestris L.) and Silver birch (Betula pendula Roth.) woodlands in a French volcanic mid-elevation area. For this purpose root biomass, root profile and fine-root architecture of competitor trees were examined in 53 mixed pine–beech and 42 birch–beech woodlands along a stand maturation gradient, using the root auger technique (0–75-cm). The total beech fine-root biomass highly correlated with aerial dimensions such as stem height and girth, whereas it moderately correlated with its age, thus indicating the effects of competition. Basic stand biometric data such as stand density and basal area had no significant effect on beech root biomass. Conversely, competition indices taking into account the vertical dimensions of competitor trees were efficient, probably due to redundancy with beech height. At similar age and height, beeches under birch had a greater belowground development than beeches under pine. Each species exhibited specific rooting pattern and plasticity of fine-root architecture along the gradients of stand maturation and competition. Beech had a heart-shaped rooting habit in both mixings, which strongly increased along stand maturation. Its fine-root system adopted a foraging strategy to respond to increasing stand competition. The Scots pine fine-root system was plate-like and showed a low morphological plasticity, thus presumably a conservative strategy. Silver birch exhibited a high biomass and a foraging capacity in the topsoil but a loose root system in the subsoil. The coexistence of pine and beech roots in the upper soil presumably leads to a high belowground competition. Beech root system becomes predominant throughout the soil profile and it adopts an efficient foraging strategy, but at the expense of its belowground development. Conversely, the niche partitioning strategy between beech and birch may explain why beech develops strongly belowground in spite of the fact that birch has a dense rooting and a competitive fine-root architecture. As a consequence, beech mid-term regeneration and development may be facilitated under birch as compared with pine.