Waterlogging responses of Sporobolus virginicus (L.) Kunth.

Summary Flooding responses in Sporobolus virginicus (L.) Kunth., a perennial C₄ grass, propagated from plants collected on the fringes of a mangrove swamp, were examined in a glasshouse study over 42 days. Flooding significantly reduced soil redox potential, induced adventitious root development, shifted resource allocation from below- to above-ground components without affecting total biomass accumulation and significantly decreased below-ground/above-ground biomass ratios. Although soil waterlogging significantly increased alcohol dehydrogenase activity (ADH) after 30 h, significant increase in central air space by 45–50% of the cross-sectional stem area eliminated root hypoxia, and ADH activity decreased to levels equivalent to drained controls after 42 days. In addition, flooded plants exhibited significantly higher carbon dioxide assimilation rates but similar relative growth rates (RGR) to drained controls. The results indicate that S. virginicus responds to water-logging by a combination of metabolic, morphological and anatomical mechanisms, which may account for its widespread distribution in coastal lagoons, estuaries and marshes.     

Phenotypic plasticity of four Chenopodiaceae species with contrasting saline–sodic tolerance in response to increased salinity–sodicity

It is unknown whether phenotypic plasticity in fitness‐related traits is associated with salinity–sodicity tolerance. This study compared growth and allocation phenotypic plasticity in two species with low salinity–sodicity tolerance (Chenopodium acuminatum and C. stenophyllum) and two species with high salinity–sodicity tolerance (Suaeda glauca and S. salsa) in a pot experiment in the Songnen grassland, China. While the species with low tolerance had higher growth and allocation plasticity than the highly tolerant species, the highly tolerant species only adjusted their growth traits and maintained higher fitness (e.g., plant height and total biomass) in response to increased soil salinity–sodicity, with low biomass allocation plasticity. Most plasticity is “apparent” plasticity (ontogenetic change), and only a few traits, for example, plant height:stem diameter ratio and root:shoot biomass ratio, represent “real” plasticity (real change in response to the environment). Our results show that phenotypic plasticity was negatively correlated with saline–sodic tolerance and could be used as an index of species sensitivity to soil salinity–sodicity.     

Increased compensatory ability of an invasive plant to above- and below-ground enemies in monocultures

Above- and below-ground enemies have prominent influence on plant invasions, and increasing evidence has shown that plant invasions are also affected by inter- or intraspecific interactions between individual plants. However, how these two factors interactively affect plant invasions has rarely been tested. Here, we examined the response of the invasive plant Alternanthera philoxeroides (Mart.) Griseb. to above- and below-ground enemies at varying plant densities in a greenhouse experiment in Wuhan, China. We found that both above- and below-ground enemies decreased the plant total and root mass at individual and population levels, but that of the two, below-ground enemies had a greater effect than above-ground enemies, and that the two guilds of enemies acted additively on the plant. However, their impacts decreased as the plant density increased, due to enhanced plant tolerance to both guilds of enemies. The increased plant tolerance may result from changes in plant resource allocation patterns, corresponding to a positive linear relationship between the ratio of fine root mass to total root mass and plant density. Given that forming dense monocultures in their new ranges is one of the most important characteristics of invasive plants, we propose that the high compensatory ability at dense monocultures may be an important mechanism underlying exotic species invasion.     

Responses of Fraxinus excelsior seedlings to grass-induced above- and below-ground competition

The competitive interactions between woody seedlings and herbaceous vegetation have received increasing interest in recent years. However, little is known about the relative contributions and underlying mechanisms of above- and below-ground competition between species. We used a novel experimental approach to assess the responses of Fraxinus excelsior seedlings to different combinations of root and shoot competition imposed by the grass Dactylis glomerata under greenhouse conditions. Seedling growth was significantly reduced by competition for soil resources, but neither biomass nor height were significantly affected by shoot competition for light. Competitive response indices based on biomass confirmed that below-ground competition was more important than above-ground competition, and indicated that root and shoot competition did not interact to influence plant growth. Fraxinus biomass allocation and seedling traits were almost all significantly affected by root competition; these responses varied depending on the trait examined. In contrast, morphological responses to shoot competition were limited. In the absence of root competition, seedlings showed a significant increase in the biomass allocated to leaves and a greater leaf area ratio in response to shoot competition. Our findings suggest that morphological modifications help to mitigate the negative effects of competition, but the expression of plasticity may be suboptimal due to resource constraints. The present study also highlights the importance of appropriate experimental controls and analysis to avoid confounding effects of experimental design and ontogeny on the interpretation of competitive responses.     

Effects of competition on root–shoot allocation in <Emphasis Type="Italic">Plantago lanceolata</Emphasis> L.: adaptive plasticity or ontogenetic drift?

We investigated how shoot and root allocation in plants responds to increasing levels of competitive stress at different levels of soil fertility. In addition, we analyzed whether different responses were due to adaptive plasticity or should be attributed to ontogenetic drift. Plantago lanceolata plants were grown during 18 weeks at five plant densities and four nutrient supply levels in pots in the greenhouse. Thereafter root and shoot biomass was measured. There were clear negative effects of increasing plant densities on plant weights revealing strong intraspecific competition. At the lower N-treatments, the proportional allocation to root mass increased with increasing competitive stress, indicating the important role of belowground competition. At the higher N-supply rate, the relationship between competitive stress and shoot to root ratio was neutral. These responses could not be attributed to ontogenetic drift, but could only be explained by assuming adaptive plasticity. It was concluded that at lower N-supplies belowground competition dominates and leads to increased allocation to roots, while at the higher N-supply competition for soil resources and light had balanced impacts on shoot and root allocation. An alternative hypothesis explaining the observed pattern is that light competition has far less pronounced impacts on root–shoot allocation than nutrient deprival.     

Allometry of Salsola collina in response to soil nutrients,water supply and population density

The allometry of greenhouse‐grown Salsola collina Pall. in response to variation in soil nutrient content, water supply and population density has been compared. The results showed that the biomass allocation was size‐dependent. Root, stem, leaf and reproductive allocation showed a ‘true’ plasticity in response to soil nutrient variation. At low soil nutrient content, plants tended to allocate more biomass to the development of reproductive organs than to stem and leaf, but root allocation was consistent due to a tradeoff between the effects of plant size and soil nutrient content. The plasticity of stem allocation and reproductive effort was ‘true’, while the plasticity of root allocation was ‘apparent’, but there was no plasticity for leaf allocation in response to soil water variation. At lower soil water content, plants tended to allocate more biomass to the stem than to development of reproductive organs. With the exception of ‘apparent’ plasticity of root allocation, no plasticity was detected in biomass allocation when population density was varied.     

Shading and the capture of localized soil nutrients: nutrient contents,carbohydrates, and root uptake kinetics of a perennial tussock grass

Summary The ability to exploit spatial and temporal heterogeneity in soil resources can be one factor important to the competitive balance of plants. Competition above-ground may limit selective plant responses to below-ground heterogeneity, since mechanisms such as root proliferation and alterations in uptake kinetics are energy-dependent processes. We studied the effect of shading on the ability of the perennial tussock grassAgropyron desertorum to take up nutrients from enriched soil microsites in two consecutive growing seasons. Roots of unshaded plants selectively increased phosphate uptake capacity in enriched soil microsites (mean increases of up to 73%), but shading eliminated this response. There were no changes in ammonium uptake capacity for roots in control and enriched patches for either shaded or unshaded plants. The 9-day shade treatments significantly reduced total nonstructural carbohydrate (TNC) concentrations for roots in 1990, but had no apparent effect on root carbohydrates in 1991 despite dramatic reductions in shoot TNC and fructan concentrations. Enrichment of the soil patches resulted in significantly greater phosphate concentrations in roots of both shaded and unshaded plants, with less dramatic differences for nitrogen and no changes in potassium concentrations. In many respects the shaded plants did surprisingly well, at least in terms of apparent nutrient acquisition. The effects of aboveground competition on nutrient demand, energy requirements, and belowground processes are discussed for plants exploiting soil resource heterogeneity.     

梓树苗木地下和地上竞争的效应研究

采用生物量计算的竞争指数和通径分析的方法,研究了3种密度的梓树苗木地下竞争和地上竞争的关系及对总竞争的影响。结果显示,梓树苗木地下生物量、地上生物量和总生物量与密度密切相关,随着密度的增加,其根、茎、叶的生物量减少,根冠比均小于1。在同一密度条件下,地上竞争指数明显大于地下竞争指数,地上竞争对总竞争的直接作用范围(0.449 3～0.973 1)明显大于地下竞争对总竞争的直接作用(0.275 6～0.773 2)。研究表明,梓树幼苗地上茎、叶的竞争在梓树苗木总的竞争中占有重要地位。     

Relative growth rate,resource allocation and root morphology in the perennial legumes, <Emphasis Type="Italic">Medicago sativa</Emphasis>, <Emphasis Type="Italic">Dorycnium rectum</Emphasis> and <Emphasis Type="Italic">D. hirsutum</Emphasis> grown under controlled conditions

Perennial pastures are needed in farming systems in southern Australia to combat environmental problems such as dryland salinity. The mediterranean climate in southern Australia imposes constraints to the growth and survival of perennial plants. The aim of this study was to compare growth rates, resource allocation and root distribution in three perennial legumes, Medicago sativa L., Dorycnium hirsutum (L.) Ser. and Dorycnium rectum (L.) Ser., to identify different plant traits and their ecological and agronomic significance. Plants were grown in 1-m deep split tubes and destructive harvests were made every 2 weeks after plant emergence for 10 weeks. Leaf area and leaf, stem and root fresh and dry weights were measured. Maximum root depth and the root distribution were also determined. Seedlings of Dorycnium were slower to emerge and had a lower relative growth rate (RGR) than M. sativa. The slower RGR was associated with a lower specific leaf area (SLA) in D. hirsutum and a lower net assimilation rate (NAR) in D. rectum. Although all species allocated a similar proportion of biomass to roots, D. rectum had a shallower root distribution and took longer to produce deep roots. The slow growth rates of Dorycnium seedlings suggest that they are more prone to establishment problems due to competition from weeds or other pastures, and because they have less access to water at greater depth during summer drought. However, D. hirsutum displayed characteristics of a plant that is adapted to stressful environments and therefore may be able to grow in conditions where other perennial legumes cannot.     

<Emphasis Type="Italic">Acer rubrum</Emphasis> (red maple) growth is negatively affected by soil from forest stands dominated by its invasive congener (<Emphasis Type="Italic">Acer platanoides</Emphasis>, Norway maple)

Invasive species continue to alter the plant communities of the eastern United States. To better understand the mechanisms and characteristics associated with invasive success, we studied competition between two Acer species. In a greenhouse, we tested (1) the effect of forest soil type (beneath an invasive and native stand) on seedling growth of the invasive Acer platanoides (Norway maple) and native A. rubrum (red maple), and the (2) effects of full (above- and below-ground) and partial inter-specific competition on species growth. We found A. rubrum growth was negatively affected by soil from the invaded stand, as it had lower above-ground (32%) and below-ground (26%) biomass, and number of leaves (20%) than in the native soil. The root:shoot resource allocations of A. platanoides depended on soil type, as it had 14% greater root:shoot mass allocation in the native soil; this ability to change root:shoot allocation may be contributing to the ecological success of the species. Widely published as having a large ecological amplitude, A. rubrum may be a useful species for ecological restoration where A. platanoides has been present, but the impacts of A. platanoides on soil functioning and subsequent plant interactions must be addressed before protocols for native reintroductions are improved and implemented.     

A Model Allocating Growth Among Leaf Proteins, Shoot Structure, and Root Biomass to Produce Balanced Activity

A model is developed that considers the allocation of carbonand nitrogen substrates to a protein compartment in the shoots,shoot structural components, and root biomass. Inclusion ofa shoot-protein compartment allows variation in shoot-specificactivity to be modelled as a function of leaf nitrogen concentration.Allocation to the biomass compartments is controlled by twopartitioning variables that are defined by explicitly usingthe balanced activity hypothesis. The model produces balancedactivity where the shoot-specific activity, as well as rootand shoot biomass, vary in response to the above-ground (lightand CO₂) and below-ground (nitrogen) environments. The predictedpatterns of both root: shoot ratio and leaf nitrogen concentrationin response to environmental resource availability are qualitativelyconsistent with general trends observed in plants. Biomass allocation, plant growth, modelling, leaf nitrogen, root: shoot ratio, balanced activity     

单叶蔷薇幼苗根系对不同潜水埋深的适应机制

地下水是影响西北地区植被分布、生长和群落演替的重要因子,通过人工装置模拟30 cm(D30)、40 cm(D40)、50 cm(D50)、60 cm(D60)、70 cm(D70)5个潜水梯度,从生长发育、根系形态、拓扑结构与分形维数以及表型可塑性四个方面来分析不同潜水埋深对单叶蔷薇幼苗的影响,力求揭示单叶蔷薇幼苗对不同水分环境的适应性策略,这将对今后开展单叶蔷薇植被恢复和保育工作具有重要价值。研究结果表明:(1)单叶蔷薇幼苗可通过增加扎根深度、总根长、根表面积、根体积、根尖数量、分支数量、地上干物质和根系干物质来应对不同潜水埋深带来的干旱胁迫,D50、D60、D70和CK处理下的幼苗还可以通过提高根冠比来适应更长久的干旱环境。(2)不同潜水埋深处理下,单叶蔷薇幼苗根系的拓扑指数基本保持在0.8-0.9之间,说明该根系属于典型的人字形分支模式,受环境影响较小。其中,短而细的密集细根(0-2 mm)构成了单叶蔷薇幼苗根系的主体。从资源分配的角度来看,该种拓扑结构相对简单、内部竞争较小、碳消耗少,有利于根系扩大土壤资源获取效率,从而保障植株生长发育的物质供需平衡,这是单叶蔷薇对环境胁迫的适应性策略。(3)适度的干旱,如50-70 cm的潜水埋深,可以促进单叶蔷薇幼苗扎根深度;而在较浅的潜水埋深(30-40 cm)环境中,单叶蔷薇幼苗能快速解除干旱,转向地上器官的生长发育,同时它通过降低垂直根系长度、增加分支和根尖数量来获取更多氧气和适应水分充足的新环境,到第75天时生长旺盛,在株高、总根长、根表面积、根体积、根尖数量、分支数量、地上干物质、根系干物质、根组织密度和分形维数10个指标上与CK组具有显著差异,说明单叶蔷薇幼苗对水分充足和严重干旱的极端环境均有较好的适应能力,表型可塑性强。     

Allometric growth, disturbance regime, and dilemmas of controlling invasive plants: a model analysis

Disturbed communities are observed to be more susceptible to invasion by exotic species, suggesting that some attributes of the invaders may interact with disturbance regime to facilitate invasion success. Alternanthera philoxeroides, endemic to South America, is an amphibious clonal weed invading worldwide. It tends to colonize disturbed habitats such as riparian zones, floodplain wetlands and agricultural areas. We developed an analytical model to explore the interactive effects of two types of physical disturbances, shoot mowing and root fragmentation, on biomass production dynamics of A. philoxeroides. The model is based on two major biological assumptions: (1) allometric growth of root (belowground) vs. shoot (aboveground) biomass and (2) exponential regrowth of shoot biomass after mowing. The model analysis revealed that the interaction among allometric growth pattern, shoot mowing frequency and root fragmentation intensity might lead to diverse plant ‘fates’. For A. philoxeroides whose root allocation decreases with growing plant size, control by shoot mowing was faced with two dilemmas. (1) Shoot regrowth can be effectively suppressed by frequent mowing. However, frequent shoot mowing led to higher biomass allocation to thick storage roots, which enhanced the potential for faster future plant growth. (2) In the context of periodic shoot mowing, individual shoot biomass converged to a stable equilibrium value which was independent of the root fragmentation intensity. However, root fragmentation resulted in higher equilibrium population shoot biomass and higher frequency of shoot mowing required for effective control. In conclusion, the interaction between allometric growth and physical disturbances may partially account for the successful invasion of A. philoxeroides; improper mechanical control practices could function as disturbances and result in exacerbated invasion.     

The representation of root processes in models addressing the responses of vegetation to global change

The representation of root activity in models is here confined to considerations of applications assessing the impacts of changes in climate or atmospheric [CO₂]. Approaches to modelling roots can be classified into four major types: models in which roots are not considered, models in which there is an interplay between only selected above-ground and below-ground processes, models in which growth allocation to all parts of the plants depends on the availability and matching of the capture of external resources, and models with explicit treatments of root growth, architecture and resource capture. All models seem effective in describing the major root activities of water and nutrient uptake, because these processes are highly correlated, particularly at large scales and with slow or equilibrium dynamics. Allocation models can be effective in providing a deeper, perhaps contrary, understanding of the dynamic underpinning to observations made only above ground. The complex and explicit treatment of roots can be achieved only in small-scale highly studied systems because of the requirements for many initialized variables to run the models.     

The space-use strategy of plants with different growth forms,in a field experiment with manipulated nutrients and light

The biomass allocation pattern in plants is known to depend on the below and above-ground resource availabilities. In a herbaceous multi-species stand, it can be expected that the effects of nutrient and light availability on plants’ general space-use strategy are fundamentally different. We hypothesized that nutrient status alters the amount of biomass produced per unit canopy volume (biomass density), but not so much the biomass vertical distribution pattern. Changes in light availability, in contrast, should affect the vertical distribution pattern of biomass but not biomass density. We were also interested in whether the effect of resource manipulation on a plant’s space-use strategy depends on its basic morphological characteristics (growth form). The results from a four-year permanent plot experiment in a species-rich grassland, with fertilization and additional illumination from mirrors applied to 40 × 40 cm plots, showed that our main hypothesis was correct. Fertilization significantly affected biomass density above as well as below-ground, while additional illumination generally did not. Light addition altered the vertical distribution pattern of above-ground biomass, which remained unaffected by the fertilizer treatment.     

The influence of abiotic stress and phenotypic plasticity on the distribution of invasive Alternanthera philoxeroides along a riparian zone

Relatively few studies have compared invasibility and species invasiveness among microhabitats within communities, synchronously. We surveyed the abundance and performance of non-native Alternanthera philoxeroides (Mart.) Griseb. (alligator weed), its co-occurring native congener, Alternanthera sessilis (L.) DC. (sessile joyweed), and other species in a wetland community along a riparian zone in southeast China to test the hypotheses that: i) degree of invasion differs between different types of microhabitats within the community; and ii) microhabitat types that differ in invasibility also differ in soil resource availability or in sediment characteristics likely to affect resource availability; iii) phenotypic plasticity of A. philoxeroides may play a key role in its adaptation to diverse habitats as can be concluded from its extremely low genetic diversity in China. The study riparian zone comprises different types of microhabitats including wet abandoned field, swamp, marsh dunes and gravel dunes. Consistent with these hypotheses, cover of A. philoxeroides was high in abandoned fields (73 ± 2.9%) and swamps (94 ± 1.3%), which had high soil nutrients and water availability. On the contrary, cover of native A. sessilis was relatively high in marsh dunes and grave dunes, which had coarse gravel surfaces, low soil nutrients and low water availability. A. philoxeroides showed greater morphological plasticity in response to habitat variation. In abiotically harsh habitats, stems had limited growth, and were prostrate with weak adventitious roots at nodes, forming thin, scattered patches. In the two richer habitats, the highly branched plants spread over the water or soil surface, supporting dense stronger leaf-bearing stems which grew vertically. The growth pattern of A. sessilis among microhabitats did not exhibit significant variations. These results suggest that morphological plasticity and microhabitat types with high soil resources may facilitate invasions of A. philoxeroides.     

Sexual dimorphism in clonal growth forms and ramet distribution patterns in <Emphasis Type="Italic">Rumex acetosella</Emphasis> (Polygonaceae)

We investigated clonal traits in the dioecious herb Rumex acetosella to characterize sexual dimorphism in clonal forms and to correlate below-ground clonal patterns and above-ground ramet distributions. We recorded creeping root length, branching patterns, ramet and clump (caespitose ramets from the same position on the root) sprouting patterns, and biomass allocations in three females and males. We also estimated the patch size of flowering ramets within a quadrat. No sexual dimorphism was detected in the frequencies of branches and flowering ramets per root length. Male plants allocated proportionally more biomass to below-ground organs. Total root length did not differ between the sexes. Females sprouted more clumps with fewer flowering ramets per root length than males, which sprouted fewer clumps with more flowering ramets, which meant that clump sprouting patterns were phalanx-like in females and guerrilla-like in males. Flowering ramets were aggregately distributed in both females and males and patch sizes were similar between sexes, indicating that the spreader propagations were not found in the guerrilla-like males. We assumed that sexual dimorphism occurred in response to physiological integration for higher reproductive effort in females.     

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     

水曲柳苗木地下竞争与地上竞争的定量研究

对3种密度进行栽培试验,利用通径分析的方法,研究了水曲柳地下竞争和地上竞争的关系及对总竞争的影响。结果表明,水曲柳苗木的地下部分生物量、地上部分生物量和总生物量与营养空间有密切关系。随着苗木空间距离增加,由生物量计算的竞争指数下降。在同一密度条件下,地下竞争指数明显大于地上竞争指数。由于地下生长与地上生长的相互作用,各竞争指数之间具有明显的相关性。但是地下竞争和地上竞争对总竞争的影响是不同的,通径分析可以定量的区分地下竞争和地上竞争的相对大小。地下竞争对总竞争的直接作用范围在0．5543～0．7426之间,明显大于地上竞争对总竞争的直接作用(0．2851～0．5282)。随着距离的增加,单株苗木的生长空间加大,地上部分的竞争作用增加,地下部分的竞争程度减弱。但是,地下根系的竞争在水曲柳苗木总的竞争中占有重要地位。