Kin recognition: Competition and cooperation in Impatiens (Balsaminaceae)

The ability to recognize kin is an important element in social behavior and can lead to the evolution of altruism. Recently, it has been shown that plants are capable of kin recognition through root interactions. Here we tested for kin recognition in a North American species of Impatiens that has a high opportunity of growing with kin and responds strongly to aboveground competition. We measured how the plants responded to the aboveground light quality cues of competition and to the presence of root neighbors and determined whether the responses depended on whether the neighbors were siblings or strangers. The study families were identified by DNA sequencing as members of the same species, provisionally identified as Impatiens pallida (hereafter I. cf. pallida). We found that I. cf. pallida plants were capable of kin recognition, but only in the presence of another plant's roots. Several traits responded to relatedness in shared pots, including increased leaf to root allocation with strangers and increased stem elongation and branchiness in response to kin, potentially indicating both increased competition toward strangers and reduced interference (cooperation) toward kin. Impatiens cf. pallida responded to both competition cues simultaneously, with the responses to the aboveground competition cue dependent on the presence of the belowground competition cue.     

植物竞争对3种移植树苗生长的影响

在植物幼苗生长过程中,总是受到包括地下根竞争在内的各种竞争影响。植物间的竞争主要同有效光辐射、水分和各种营养相关。当外来植物侵入森林群落时,可能受到群落中其它植物竞争的影响。该文通过移植尾叶桉(Eucalyptus urophylla)、大叶相思(Acacia auriculaeformis)两种外来种和本地种荷木(Schima superba)幼苗,挖沟排除根竞争和砍树创造林窗来排除地上竞争的野外实验,研究植物竞争对幼苗生长的影响。有根竞争时,荷木、尾叶桉和大叶相思幼苗的生物量和净初级生产力均小于没有根竞争时,可见地下根竞争对3种幼苗生长有抑制性影响。尾叶桉、大叶相思和荷木3种幼苗受到的总竞争强度分别是0.357 9、0.338 3和0.198 9,受到的地下根竞争强度则分别是0.104 3、0.053 04和0.118 8,受到的地上竞争强度则分别是0.285 1、0.277 0和0.090 85。尾叶桉和大叶相思两种幼苗受到的总竞争强度间的差异不显著,但都显著大于荷木;3种幼苗受到的地上竞争强度间的差异同总竞争强度情况相似;尾叶桉和荷木两种幼苗受到的地下根竞争强度间的差异不显著,但都显著大于大叶相思。地上竞争对阳性树种尾叶桉和大叶相思两种幼苗的生长影响大,而地下根竞争则对耐阴性强的荷木幼苗影响大。尾叶桉和大叶相思两种外来种幼苗受到的总竞争强度均大于本地种荷木幼苗,这反映了这两种外来树种侵入次生林这样的群落受到竞争影响大。     

Neighbor species differentially alter resistance phenotypes in Plantago

In this study, we investigated how neighbors (i.e., competitors) altered resistance phenotypes, namely plant size and levels of secondary compounds (iridoid glycosides), of individual plants and specifically tested whether neighbor identity mattered. We conducted a greenhouse experiment with Plantago lanceolata and Plantago major (Plantaginaceae) in which each species served as focal plants as well as neighbors in a factorial design. In addition, we harvested plants six and nine weeks after transplantation to test whether effects changed as plants grew. In both species, competition reduced plant size, and this effect increased over time. Plantago lanceolata neighbors suppressed growth of both focal plant species more than P. major neighbors. Effects of competition on levels of secondary compounds were more complex. Concentrations of iridoid glycosides were increased by competition in both species at harvest one. By the second harvest, an effect of competition on iridoid glycosides was found only in P. major. Neighbor identity influenced levels of iridoid glycosides in P. lanceolata at harvest one; concentrations were higher in plants grown with P. lanceolata neighbors than in plants grown with P. major neighbors. We also tested whether there was a trade-off between growth (biomass) and defense (levels of iridoid glycosides). Biomass and iridoid glycoside content were significantly correlated only in plants grown with competition and harvested at nine weeks, and this relationship was positive in both species, indicating that there was no trade-off between growth and defense. This study suggests that neighbor identity could play an important role in interspecific interactions, including the interactions of plants with other trophic levels.     

The relative importance of above- versus belowground competition for tree growth during early succession of a tropical moist forest

Competition between neighboring plants plays a major role in the population dynamics of tree species in the early phases of humid tropical forest succession. We evaluated the relative importance of above- versus below-ground competition during the first years of old-field succession on soil with low fertility in Southern Mexico, using the premise that competition for light is size-asymmetric, unlike competition for nutrients. Plant growth is thus expected to be disproportionally impeded by larger neighbors. We studied how growth and survival of 3.5–5.5 m tall saplings of Cecropia peltata and Trichospermum mexicanum, two pioneer species that dominate the secondary forests in the study region, varied with the abundance and size of neighboring trees in 1–2 year old secondary vegetation. We found that local neighborhood basal area varied 10-fold (3 to 30 cm² m^-2) and explained most of the variation in diameter and height growth of the target saplings. Most growth variables were strongly affected by the neighbors bigger than the focal trees with no significant additive effect of the smaller neighbors, indicating asymmetric competition. Smaller neighbors did have a small but significant additive effect on the diameter growth of Cecropia saplings and stem slenderness of Trichospermum saplings. We conclude that competition for light was more important than belowground competition in this initial phase of moist tropical forest successional, despite the low soil fertility.     

Ecophysiological Responses to Neighbor Removal in an Old-Field and a Prairie in Northeastern Kansas,USA

An ecophysiological approach was used to determine if competition can be detected among plants in a recently abandoned old-field and in a native tallgrass prairie in northeastern Kansas. In situ photosynthetic parameters and water potentials (Ψ) of target plants were measured 1-2 d after neighbor (intra- and interspecific) removal as well as 1-4 weeks later, and compared with values for plants with neighbors. Only two of the six study species (four old-field and two prairie species) responded to removal of neighboring plants, and only after several weeks had elapsed. Net photosynthetic rates (P_N) and stomatal conductances (g_s) of Ambrosia trifida in an old-field increased after removal of both intra- and interspecific neighbors. For Apocynum cannabinum, another old-field species, P_N of target plants without neighbors was significantly higher than that of target plants with neighbors. For both these species, values of Ψ were not different between target plants with and without neighbors, suggesting that increased availability of nutrients may have been responsible for the observed ecophysiological responses. Though numerous past studies indicate that competition is a major factor influencing plants in old-field and in prairie communities, the experimental approach used in this study revealed that neighbor removal had only limited effects on ecophysiology of the target plants in either community. This revised version was published online in June 2006 with corrections to the Cover Date.     

Physiological regulation and functional significance of shade avoidance responses to neighbors

Plants growing in dense vegetations compete with their neighbors for resources such as water, nutrients and light. The competition for light has been particularly well studied, both for its fitness consequences as well as the adaptive behaviors that plants display to win the battle for light interception. Aboveground, plants detect their competitors through photosensory cues, notably the red:far-red light ratio (R:FR). The R:FR is a very reliable indicator of future competition as it decreases in a plant-specific manner through red light absorption for photosynthesis and is sensed with the phytochrome photoreceptors. In addition, also blue light depletion is perceived for neighbor detection. As a response to these light signals plants display a suite of phenotypic traits defined as the shade avoidance syndrome (SAS). The SAS helps to position the photosynthesizing leaves in the higher zones of a canopy where light conditions are more favorable. In this review we will discuss the physiological control mechanisms through which the photosensory signals are transduced into the adaptive phenotypic responses that make up the SAS. Using this mechanistic knowledge as a starting point, we will discuss how the SAS functions in the context of the complex multi-facetted environments, which plants usually grow in.Key words: competition, shade avoidance, hormones, cell wall, adaptive plasticity, photoreceptor, light     

The effects of neighbors on the growth and survival of shrub seedlings following fire

Where plant species vie for limited resources, disturbances might preclude competition by releasing a flush of nutrients, or by reducing biomass and thereby diminishing the consumption of resources. However, if new seedlings colonize in clumps, they may still deplete resources within the local aggregations, which may then reduce their growth and survivorship. We investigated competition among seedlings in a burned area by examining the relationship between the performance of newly established shrub seedlings of Ceanothus impressus and (1) the proximity and (2) the identity of their near neighbors. We also investigated the relationship between neighbor proximity and the availability of water. Both survivorship and growth of C. impressus were positively associated with increasing distance to near neighbors, in a manner consistent with resource competition. The availability of water (as determined by pre-dawn xylem pressure potentials) tended to be greater when neighbors were farther away, providing evidence that water was a resource for which plants were competing. This conclusion is reinforced by the finding that the effects of neighbors were stronger in drought years, suggesting that yearly variation in the availability of an important resource (water) can affect the strength of competitive interactions. This suggests that after disturbances, when some resources are apparently abundant on a large scale, competition may be important in determining the small scale patterns of seedling growth and survival.     

Plant competition and exclusion with optimizing individuals

Most models of plant competition represent competition as taking place between species when realistically competition takes place between individuals. We model individual plants as optimally choosing biomass in order to maximize net energy that is directed into reproduction. Competition is for access to light and a plant that grows more biomass adds to the leaf area index, creating negative feedback in the form of more self shading and shading of its neighbors. In each period and for given species densities, simultaneous maximization by all plants yields an equilibrium characterized by optimum biomasses. Between periods the net energies plants obtain are used to update the densities, and if densities change the equilibrium changes in the subsequent period. A steady state is attained when all plants have net energies that just allow for replacement. Four main predictions of the resource-ratio theory of competition are obtained, providing behavioral underpinnings for species level models. However, if individual plant parameters are not identical across species, then the predictions do not follow. The optimization framework yields many other predictions, including how specific leaf areas and resource stress impact biomass and leaf area indices.     

林窗中植物竞争强度随林窗发育的变化

 通过野外移植外来种尾叶桉(Eucalyptus urophylla)、大叶相思(Acacia auriculaef ormis)和本地种荷木(Schima superba) 3种幼苗,挖沟排除根竞争和砍树创造林窗排除地上竞争的实验,来测定林窗形成后植物地下竞争和地上竞争强度的变化以及这些竞争对3种幼苗生长的影响。在林窗内,尾叶桉、大叶相思和荷木受到的地下竞争强度和地上竞争强度均呈现出随林窗年龄的增长而逐渐增大的趋势。外来种尾叶桉和大叶相思受到的地上竞争强度大于本地种荷木受到的地上竞争强度,而荷木受到的地下竞争强度大于两种外来种受到的地下竞争强度。在只有地下竞争和地上竞争时,这3种幼苗的相对生长速率均小于没有竞争时,这反映了地下竞争和地上竞争对3种幼苗的生长有抑制性影响,而且这种影响在本地种和外来种间存在差异。     

Plant size and spatial pattern in a natural population of Myosotis micrantha

Abstract. We examined spatial distributions and plant sizes along a transect through a natural population of a winter annual, Myosotis micrantha. A size hierarchy existed, as indicated by high values of Gini coefficients of inequality for plant mass and correlated measures. Plants with no immediate conspecific neighbors were larger than plants with one or more near neighbors, suggesting that competition from near neighbors depressed plant size. However, there was strong positive spatial autocorrelation in plant size: large plants were associated with large neighbors and small ones with small neighbors. Plant size was also positively correlated with the combined biomass of near neighbors. The population formed a two-phase mosaic of patches of relatively large plants alternating with patches of smaller plants. The data suggest that individual plants compete with conspecifics, but the effects of competition are symmetrical. The most likely explanations for this spatially structured size hierarchy are variation in plant density, patchy distribution of resources, or a combination of the two.     

Associational resistance mediates interacting effects of herbivores and competitors on fireweed performance

Neighboring plants may have dual roles in relation to focal plants, being both competitors, and mediators of associational resistance (AR), or associational susceptibility (AS) to herbivores. Even though these two roles are played out simultaneously in nature, they have traditionally been studied separately. We conducted a two-by-two factorial experiment by manipulating the occurrence of neighbors (meadowsweet, Filipendula ulmaria) and shared mammal herbivores (moose, Alces alces) to investigate how the two simultaneously affect the performance of focal fireweed (Chamerion angustifolium) plants.We found that interactions between mammal herbivory and neighbors had significant effects on fireweed performance. These effects were explained by two counteracting mechanisms. First, when mammal herbivores were present, fireweed experienced AR such that plants growing outside of meadowsweet patches had an almost tripled risk of being browsed compared to plants growing among meadowsweet, resulting in reduced fireweed reproduction and growth. Second, when mammals were excluded, the association with meadowsweet had a direct negative effect on fireweed growth rates – mainly due to competition for light. Additionally, for the first time, we show that association with neighbors affected mammal-induced branching in fireweed.This study demonstrates that AR mediated by mammal herbivores can flip plant–plant interactions from competitive (−) to facilitative (+), and affect plant architecture. As AR and competition are probably simultaneously co-occurring forces, interactions between the two are likely common and thus of fundamental importance for populations and communities.     

Mechanisms determining the degree of size asymmetry in competition among plants

When plants are competing, larger individuals often obtain a disproportionate share of the contested resources and suppress the growth of their smaller neighbors, a phenomenon called size-asymmetric competition. We review what is known about the mechanisms that give rise to and modify the degree of size asymmetry in competition among plants, and attempt to clarify some of the confusion in the literature on size asymmetry. We broadly distinguish between mechanisms determined primarily by characteristics of contested resource from those that are influenced by the growth and behavior of the plants themselves. To generate size asymmetric resource competition, a resource must be “pre-emptable.” Because of its directionality, light is the primary, but perhaps not the only, example of a pre-emptable resource. The available data suggest that competition for mineral nutrients is often size symmetric (i.e., contested resources are divided in proportion to competitor sizes), but the potential role of patchily and/or episodically supplied nutrients in causing size asymmetry is largely unexplored. Virtually nothing is known about the size symmetry of competition for water. Plasticity in morphology and physiology acts to reduce the degree of size asymmetry in competition. We argue that an allometric perspective on growth, allocation, resource uptake, and resource utilization can help us understand and quantify the mechanisms through which plants compete. Received: 17 February 1997 / Accepted: 8 October 1997     

Complex network analysis reveals novel essential properties of competition among individuals in an even-aged plant population

Little is known about the network structure of competition in large populations of plants, despite the importance of such knowledge for understanding population dynamics. In this study, we used complex network analysis to examine temporal changes in the network structure of competition in an even-aged multi-individual stand of the Sakhalin fir Abies sachalinensis in Hokkaido, Japan. Using census data, which were measured over 30 years (1948–1978; seedlings were planted in 1929), on the sizes and locations of these plants, we regarded a plant as a node and competition between plants as a link. We then introduced two indices, the binary and weighted out-degrees (BO and WO, respectively), to interpret complicated plant interactions. The BO of a plant represents the number of links from the target plant to its neighbors, and the WO is the total strength of competition from the target plant to its neighbors. The analysis showed that the distributions of BO and WO were heavy-tailed in all years and that large plants had large BO and WO. These results suggest that only a few (i.e., large) plants have a very large impact on the growth and survival of a much larger number of neighboring plants and thus on population dynamics, whereas most of the others (i.e., small and medium-sized plants) have only a small impact on a few neighbors. By introducing binary and weighted connectivities (BC’ and WC’, respectively), we were able to identify the size classes of neighbors with which the target plant preferentially and strongly competed. The BC’ and WC’ results showed that large plants competed preferentially and more strongly with other large plants in 1948, but they competed more strongly with small plants after 1963. These results clarify targets of the very large impact of large plants, as shown by the results of BO and WO: the impact was exerted on the growth and survival of other large plants in 1948, whereas the impact was exerted on those of small plants after 1963. Our study demonstrates that the statistical properties of the competition network structure, which have been largely ignored in plant competition research, are important for understanding plant population dynamics.     

Intraspecific interactions in the annual legume Medicago minima are shaped by both genetic variation for competitive ability and reduced competition among kin

Documenting if plants exhibit kin competition avoidance in intraspecific plant interactions is relevant both to improve crop growth, and to understand diversity and composition in natural plant communities. However, a number of confounding mechanisms complicates detecting kin competition avoidance from experiments comparing plants growing with kin and non-kin neighbors. We conducted complementary greenhouse experiments using genotypes from four populations of the annual Medicago minima, which in a previous study showed higher survival when interacting with kin relative to non-kin. We show that genotypes vary in kin competition avoidance, and in competitive ability, but find no indication of complementary resource use. Importantly, from our first experiment of root growth behavior, we know that some genotypes exhibit kin competition avoidance. Yet, the variation in competitive ability we find in our second experiment, where plants grow in mini communities together with either kin or unrelated genotypes, can alone explain the variation we observe in growth and biomass among communities. In our case, the genotypes with highest competitive ability were also those that showed kin competition avoidance. This confounding effect obscured the disentangling of mechanisms underlying difference in growth between kin and non-kin interactions. When silencing root exudates by adding activated carbon to a subset of our genotype combinations, we found increased size asymmetry of plants grown together, and mostly in kin communities. This suggests that plants recognize the identity of neighbors via root exudates, and compete less with neighbors recognized as kin. To detect kin competition avoidance we suggest designing experiments that pair unrelated genotypes with similar competitive abilities. Such design, combined with silencing root exudates would be powerful to detect whether plants show kin competition avoidance or not.     

Differing selection in alternative competitive environments: shade-avoidance responses and germination timing

Under competitive conditions, stem elongation in plants is thought to enhance fitness by increasing light interception. However, the onset of competition should vary with the species of competitor due to interspecific differences in timing of emergence and plant growth form. The fitness benefits of elongation may therefore depend on the timing of this plastic response. Phenotypic selection analyses and path analysis were used to evaluate selection acting on stem elongation at early and late life-history stages and the combination of germination timing and elongation in an annual plant. Velvetleaf (Abutilon theophrasti) were raised in one of three environments experienced by natural populations (cornfields; soybean fields; and disturbed, weedy sites). Due to the rapid growth rate and high density of plants in disturbed areas, selection to increase seedling-stage elongation was expected in weedy sites. Due to the wide spacing of crop plants, competition for light is initially low in cultivated fields, but intensifies as the season progresses. Selection for increased elongation at later nodes was expected in soybean fields because velvetleaf can often overtop soy and thereby increase leaf exposure. In contrast, selection against late elongation was expected in cornfields because velvetleaf are incapable of overtopping corn. Individuals that elongate would experience the carbon cost of allocating to structural tissue, but fail to experience a carbon return through increased light interception. The phenotypic selection analyses were consistent with these predictions and therefore support the role of stem elongation as an adaptation to interspecific competition. Selection also acted on the combination of germination timing and elongation. In the weedy environment, early emergence in conjunction with enhanced stem elongation conveyed the highest fitness. Reduced elongation was favored among individuals that emerged late, potentially because these individuals were unable to overtop neighbors. The results of this study demonstrate that the timing of stem elongation strongly affects competitive success. Environments that differ in the timing of competition for light select for elongation at different life-history stages, and this selection depends on the timing of emergence.     

Plant adaptation to dynamically changing environment: the shade avoidance response

The success of competitive interactions between plants determines the chance of survival of individuals and eventually of whole plant species. Shade-tolerant plants have adapted their photosynthesis to function optimally under low-light conditions. These plants are therefore capable of long-term survival under a canopy shade. In contrast, shade-avoiding plants adapt their growth to perceive maximum sunlight and therefore rapidly dominate gaps in a canopy. Daylight contains roughly equal proportions of red and far-red light, but within vegetation that ratio is lowered as a result of red absorption by photosynthetic pigments. This light quality change is perceived through the phytochrome system as an unambiguous signal of the proximity of neighbors resulting in a suite of developmental responses (termed the shade avoidance response) that, when successful, result in the overgrowth of those neighbors. Shoot elongation induced by low red/far-red light may confer high relative fitness in natural dense communities. However, since elongation is often achieved at the expense of leaf and root growth, shade avoidance may lead to reduction in crop plant productivity. Over the past decade, major progresses have been achieved in the understanding of the molecular basis of shade avoidance. However, uncovering the mechanisms underpinning plant response and adaptation to changes in the ratio of red to far-red light is key to design new strategies to precise modulate shade avoidance in time and space without impairing the overall crop ability to compete for light.     

EFFECTS OF COMPETITION AND DISTURBANCE ON THE RESPROUTING PERFORMANCE OF THE MEDITERRANEAN SHRUB ERICA MULTIFLORA L. (ERICACEAE)

Two field experiments were designed to evaluate the importance of competition, fire, repeated disturbance, and their interactions on the vegetative and reproductive performance of the Mediterranean shrub Erica multiflora over a 2.5-yr period. In a burn experiment, fire was applied to the ground-level stumps of previously clipped 13-yr-old plants with a propane torch and competition was diminished by removal of neighboring plants. Fire resulted in a reduction of sprout vigor and biomass of flowers; mature neighbors also reduced E. multiflora sprout vigor and flowering. The interaction between fire and competition was nonsignificant. In a stand burned by a wildfire we studied the effects of regenerating neighbors on target plants by removing all neighbors or only Quercus coccifera, the most dominant species in the burned stand. In this stand we also simulated herbivory by repeatedly clipping the sprouts of E. multiflora. Regenerating neighbors did not affect target plant sprout vigor after the wildfire, but did cause a decrease in the biomass of flowers per plant. Survival decreased after repeated clipping but was not affected by neighborhood treatment. The results suggest that the importance of competition on resprouting vigor was temporally variable. Variables related to plant size rather than species determined competitive superiority: resprouting neighbors did not affect resprouting performance of target plants, but mature neighbors did. In nature, fire may directly reduce vegetative and reproductive biomass by the heating effect. But it may have an indirect positive effect on biomass, by reducing competition among plants. Frequent disturbances that removed aboveground biomass of E. multiflora had a detrimental effect on target plant survival independent of neighborhood effect.     

Competition intensity along a productivity gradient in a low-diversity grassland

Competition among plants often shifts from roots to shoots as productivity increases and species composition changes. We examined competition in an old field with low diversity to test whether this shift occurred along a productivity gradient without species turnover. Forty plots received one of four nitrogen treatments (0, 5, or 15 g added N m-2 yr-1 or 400 g m-2 yr-1 of sawdust added to immobilize N) annually for 5 yr. All N levels were dominated by the perennial grasses Agropyron cristatum and Bromus inermis. Transplants of Agropyron were grown with all neighbors, roots of neighbors, or no neighbors present to measure total, root, and shoot competition. Transplant growth was 22%-165% higher in subplots without neighbors present, which indicates that competition occurred. Competition from neighbor roots was primarily responsible for suppression of transplant growth over the entire productivity gradient. In contrast to previous field experiments that found either an increase in total competition intensity or a shift from root to shoot competition with increasing productivity, we found neither. Increases in total competition intensity or shifts from root to shoot competition found along other gradients may be caused by changes in species composition and not by increased resources or neighbor biomass. These results suggest that different competitive mechanisms may operate in low-diversity vegetation than in more diverse natural vegetation.     

Investigating the relationship between neighbor root biomass and belowground competition: field evidence for symmetric competition belowground

Little is known about how small-scale variation in neighbor biomass can influence the strength of root competition experienced by an individual plant. In this study, modified root exclusion tubes were used to vary the accessibility of the soil space surrounding Amaranthus retroflexus target plants to the neighboring plants. A gradient of root accessibility was created by drilling varying numbers of holes into standard root exclusion tubes, made of 15 cm diameter PVC pipe. Belowground competitive intensity, defined as biomass reduction due to root interactions alone, relative to growth in the absence of neighbors, was then measured along the resulting gradient of neighbor root densities. At low neighbor root abundances the strength of belowground competition was proportional to neighbor root biomass, consistent with prior evidence that belowground competition is symmetric. If belowground competition were asymmetric, neighbor roots should have had little effect on target plants when they are rare relative to those of the target plant. At higher neighbor root abundances, belowground competitive intensity should increase rapidly. The strong relationship found between neighbor root biomass and belowground competitive intensity suggests relatively small variations in root biomass could lead to large variations in belowground competition. Reduced belowground competition in areas with low root biomass could have important implications for the establishment and growth of poor belowground competitors, suggesting a mechanism by which species coexistence may occur despite extremely intense root competition.     

EVOLUTION OF BRASSICA RAPA L. (CRUCIFERAE) POPULATIONS IN INTRA- AND INTERSPECIFIC COMPETITION

Populations of Brassica rapa were grown for three generations in each of two environments: intraspecific competition, with four surrounding Brassica rapa neighbors per pot, and interspecific competition, with two Raphanus sativus neighbors per pot. In each environment, the largest (by flower number) 10% of the plants were outcrossed and provided seeds for the next generation. As a control, a randomly chosen 10% of the plants in each environment were outcrossed to produce seed for the next generation. Each of these four treatments, the selected lines in intra- and interspecific competition and the corresponding control lines, was maintained for three generations. After a single generation of growth in a common, no-competition environment, replicate plants from each treatment were grown with no competition and with intra- and interspecific competition for determination of growth responses. After two generations of selection, flower number in the intraspecific-selection line had increased by more than 50% over that in the control line and by more than 19% over that under interspecific selection. After a common-environment generation, plants from the intraspecific-selection line were shown to have significantly faster growth in height and flower number as seedlings. Plants in the interspecific-selection line showed similar but nonsignificant trends. No differences in seed mass, emergence time, or photosynthetic rate were found between control and selected lines in either intra- or interspecific competition. Some differences between control and selected lines were noted in biomass allocation related to differences in phenology. The results demonstrate that performance in competitive environments can evolve through changes in plant development but that rates of evolution will differ in intra- and interspecific competition.