Plant competition for light analyzed with a multispecies canopy model

Summary Competition for light among species in a mixed canopy can be assessed quantitatively by a simulation model which evaluates the importance of different morphological and photosynthetic characteristics of each species. A model was developed that simulates how the foliage of all species attenuate radiation in the canopy and how much radiation is received by foliage of each species. The model can account for different kinds of foliage (leaf blades, stems, etc.) for each species. The photosynthesis and transpiration for sunlit and shaded foliage of each species is also computed for different layers in the canopy. The model is an extension of previously described single-species canopy photosynthesis simulation models. Model predictions of the fraction of foliage sunlit and interception of light by sunlit and shaded foliage for monoculture and mixed canopies of wheat (Triticum aestivum) and wild oat (Avena fatua) in the field compared very well with measured values. The model was used to calculate light interception and canopy photosynthesis for both species of wheat/wild oat mixtures grown under normal solar and enhanced ultraviolet-B (290–320 nm) radiation (UV-B) in a glasshouse experiment with no root competition. In these experiments, measurements showed that the mixtures receiving enhanced UV-B radiation had a greater proportion of the total foliage area composed of wheat compared to mixtures in the control treatments. The difference in species foliage area and its position in the canopy resulted in a calculated increase in the portion of total canopy radiation interception and photosynthesis by wheat. This, in turn, is consistent with greater canopy biomass of wheat reported in canopies irradiated with supplemental UV-B.     

Early-season effects of supplemented solar UV-B radiation on seedling emergence, canopy structure, simulated stand photosynthesis and competition for light

Mixtures and monocultures of wheat (Triticum aestivum) and wild oat (Avena fatua), a common weedy competitor of wheat, were exposed to enhanced solar UV-B radiation simulating a 20% reduction in stratospheric ozone to assess the timing and seasonal development of the UV-B effects on light competition in these species. Results from two years of field study revealed that UV-B enhancement had no detectable effect on the magnitude or timing of seedling emergence in either species. End-of-season measurements showed significant UV-B inhibition of leaf insertion height in wild oat in mixture and monoculture in the second year (irrigated year) but not in the first year (drought year). Leaf insertion height of wheat was not affected by UV-B in either year. The UV-B treatment had no detectable effect on monoculture or total (combined species) mixture LAI but did significantly increase (5–7%) the fractional contribution of wheat to the mixture LAI after four weeks of growth in both years. In addition, the UV-B treatment had subtle effects on LAI height profiles with early season mixtures showing significant reductions in wild oat LAI in lower canopy layers in both years while midseason Year 2 mixtures showed significant reductions in wild oat LAI in upper canopy layers. The changes in canopy structure were found to significantly increase (6–7%) the proportional simulated clear sky canopy photosynthesis and light interception of wheat in mixture. These findings, and others, indicate that the effects of UV-B enhancement on competition are realized very early in canopy development and provide additional support for the hypothesis that UV-B enhancement may shift the balance of competition between these species indirectly by altering competitive interactions for light.     

Plant competition for light analyzed with a multispecies canopy model

Summary A multispecies canopy photosynthesis simulation model was used to examine the importance of canopy structure in influencing light interception and carbon gain in mixed and pure stands of wheat (Triticum aestivum L.) and wild oat (Avena fatua L.), a common weedy competitor of wheat. In the mixtures, the fraction of the simulated canopy photosynthesis contributed by wheat was found to decline during the growing season and this decline was closely related to reductions in the amount of leaf area in upper canopy layers. For both species in mixture and in monoculture, simulated photosynthesis was greatest in the middle or upper-middle canopy layers and sensitivity analyses revealed that canopy photosynthesis was most sensitive to changes in leaf area and leaf inclination in these layers. Changes in LAI and leaf inclination affected canopy carbon gain differently for mixtures and monocultures, but the responses were not the same for the two species. Results from simulations where the structural characteristics of the two species were substituted indicated that species differences in leaf inclination, sheath area and the fraction of leaf area alive were of minor consequence compared with the differences in total leaf area in influencing relative canopy carbon gain in mixtures. Competition for light in these species mixtures appears to be influenced most by differences in the positioning of leaf area in upper canopy layers which determines, to a great extent, the amount of light intercepted.     

A simulation model of Avena fatua L. (wild-oat) growth and development

An eco-physiological simulation model of the growth and development of Avena fatua was parameterised and tested. The model simulates growth ofA. fatua, in kg dry matter ha^-1 day^-1 from sowing to maturity as a function of irradiance, temperature and various species characteristics. Parameter values were derived from the literature and from field experiments, including both autumn and spring sowings of A. fatua over three years at two sites in southern England. With two exceptions, a single set of parameter values was sufficient to accurately simulate the emergence, growth and development of both autumn and spring cohorts over all years and sites. The two exceptions were the result of differences between autumn and spring cohorts of A. fatua in the rate of early leaf area growth and in the relationship between specific leaf area and developmental stage.     

The effects of herbicides, applied atone and in sequence, on the control of wild-oats (Avena fatua) and broad-leaved weeds, and on yield of winter wheat

Experiments were done to examine the effects of controlling wild-oats and autumn-germinating broad-leaved weeds in winter wheat, early in winter or late in spring. The herbicides used were barban (winter), chlortoluron or isoproturon (winter), and benzoylprop-ethyl, at the recommended doses and at half doses. Sequential treatments of two herbicides at half doses were also examined. All treatments were given a routine broad-leaved herbicide treatment in spring. Yields of wheat were influenced more by the time of weed removal than by the degree of control achieved. Grain yields at three sites with dense autumn broad-leaved weed populations were greatest following the use of chlortoluron or isoproturon. At three other sites with moderate to dense wild-oat populations (60 to 240 plants/m¹), the use of barban at the crop three-leaf stage gave larger yields than benzoylprop-ethyl in late spring at the early stem elongation stage of the crop. Seed formation from surviving A. fatua was similar with both wild-oat herbicides. The treatment which reduced seed production of A. fatua and maintained crop yield most consistently was barban followed by benzoylprop-ethyl, each at half the normal recommended dose.     

First Quantitative Study of Rove Beetles in Oklahoma Winter Wheat Fields

Adult rove beetles (Staphylinidae) were sampled every 7–14 days from one winter wheat field located in each of the four major wheat growing regions of Oklahoma during the 1999–2000 and 2000–2001 growing seasons. The number of cereal aphids per tiller, wheat plant growth stage, and wheat tiller density also were estimated. A total of 12 genera representing 13 species of beetles were collected from the field. The density of rove beetles was generally low, ranging from 0.003 beetles per m² in fall to 0.106 beetles per m² in spring. Rove beetle communities differed among seasons. After accounting for the effect of season, there was no statistically significant association between rove beetle community structure and field location, aphid density, wheat plant growth stage, or wheat plant density. Most rove beetle species showed no association with a particular season, however, Aleochara notula Erichson, Lathrobium sp., and Oxypoda sp. were present predominantly in fall, while Bisnius inquitus Erichson was associated with winter. Oxypoda sp. was the most abundant rove beetle in winter wheat fields in spring and was relatively abundant in winter, but was not collected from wheat fields in fall. Tachyporus jocosus Say was present in wheat fields during all seasons. T. jocosus was the most abundant rove beetle species in the winter wheat fields in fall and winter and was the second most abundant species during spring.     

麦秸与稻秸浸提液对油菜及2种靶标杂草种苗生长影响的差异性研究

采用室内培养皿生物测定的方法,测试不同质量浓度（0、0.02、0.03、0.05、0.08、0.10 g·mL^-1）麦秸、稻秸浸提液对油菜及油菜田2种常见靶标杂草野燕麦（Avena fatua Linn）、播娘蒿 ［Descurainia sophia（L.）Schur］种子萌发及幼苗生长的影响。结果表明,在供试质量浓度下,麦秸与稻秸浸提液对3种受体材料种苗生长的影响存在显著差异,二者对油菜种苗生长安全的最高质量浓度分别为0.08 g·mL^-1和0.05 g·mL^-1,可以用来开发研制油菜田肥料;在对油菜安全的质量浓度下,麦秸浸提液对野燕麦种苗生长无抑制作用;而稻秸浸提液对野燕麦种苗生长的抑制作用明显,可以用来开发研制油菜田防除野燕麦的生物抑草肥;在油菜的安全的质量浓度下,麦秸、稻秸浸提液对播娘蒿种苗生长均有很强的抑制作用,且麦秸的抑制作用强于稻秸,具有开发研制油菜田治理播娘蒿的生物抑草肥料的应用前景。     

ENVIRONMENTAL CONTROL OF SEED GERMINATION IN THLASPI ARVENSE (CRUCIFERAE)

The effect of environmental conditions during storage and imbibition on germination was investigated in field pennycress (Thlaspi arvense L.), a weed species that can behave as a winter or a summer annual. Freshly harvested seeds of an inbred line with a cold requirement for flowering exhibited primary dormancy that was rapidly lost following 1 month of afterripening in a dry state. Nondormant seeds were positively photoblastic. The light effect was mediated through phytochrome since germination was promoted by red light and inhibited by far red light. Seedling emergence was also inhibited by light filtered through a canopy of wheat leaves. Germination of field pennycress seeds was considerably more sensitive to moisture stress than two sympatric species, wild oat (Avena fatua L.) and wheat (Triticum aestivum L., cv. ERA). Seeds of the latter two species were chosen in order to compare the effect of water potential on germination in field pennycress with that in sympatric species. It was concluded that the major environmental factor limiting nondormant field pennycress seeds on the soil surface was water availability. Imbibition of fully afterripened seeds at low temperatures (6 C) induced a deep secondary dormancy. In contrast to primary dormancy, cold-induced dormancy was not alleviated by red light, alternating temperatures (21/5 C), or 2 months of dry storage at 6, 15, or 35 C. However, exogenous gibberellin A₃ or 24 weeks of dry storage resulted in germination in cold-induced dormant seeds. Secondary dormancy was not observed in fully afterripened seeds that were preincubated at 21 C for 1 or 2 days prior to the cold treatment. These results may explain the failure in field experiments to observe the cold-induced secondary dormancy that limits spring emergence in other winter annuals (J. Baskin, C. Baskin, Weed Res. 1979 19: 285–292).     

Positive effects of wheat variety mixtures on aboveground arthropods are weak and variable

Although modern agriculture generally relies on homogeneous varieties that are usually grown in pure stands, crop variety mixtures have been used for a long time, notably to improve resistance to fungal diseases. A growing number of studies suggest that intraspecific plant diversity may also enhance the abundance and diversity of wild species and thereby some ecosystem services such as biological control by natural predators. However, positive effects of the genetic diversity of plant species on the diversity of associated communities have mostly been documented in natural systems, with only a handful of studies targeting crop species in agroecosystems. Here, we investigated the ecological effects of the number of winter wheat varieties (Triticum aestivum) on aboveground arthropods and particularly predatory species. We manipulated the number of wheat varieties (1, 2, 4 or 8) in 120 plots (80 m² each) to examine how wheat diversity and stand characteristics impact communities of three dominant aboveground arthropod groups that include many predatory species: ground beetles, rove beetles and spiders. The number of wheat varieties had a weak, but positive effect on predator abundance, notably spider abundance. In contrast, wheat functional diversity, as assessed by the number of wheat functional groups, was only negatively related to the diversity of spiders. Among wheat stand characteristics, the variance in plant height, wheat biomass and the Green Area Index were weakly correlated with ground beetle, rove beetle and predatory diversity, respectively. The Green Area Index was also weakly correlated with ground beetle abundance. Our study suggests that wheat variety mixtures have variable and limited effects on aboveground arthropods and probably low effectiveness to enhance biological control, but these results should be further tested under low-input agriculture in real fields.     

Population biology of avena. VI. The role of genetic polymorphisms in the outcome of interspecific competition

Three different series of population samples of two Avena species cooccurring in California were grown for an analysis of the role of genetic variation in interspecies competition: I, samples from mixed fatua-barbata sites in nature, grown in mixed stands; II, samples from pure sites and grown in pure stands, and III, the same sites as in II but grown in mixed stands. Four macroenvironments and four densities were used giving a total of sixteen entries for each genetic/competitive unit in order to measure both mean and variance of survival and reproductive rates as fitness characters. Sites used in each series included low versus high levels of genetic polymorphism within each species. In general, high polymorphism favored A. fatua in competition with monomorphic A. barbata, and high polymorphism in A. barbata allowed it to compete better with monomorphic A. fatua observation fits well into the pattern of reduced polymorphism in natural mixed stands. Mean performance of polymorphic mixed stands was not consistently higher than the monomorphic combinations or pure stands but the greater relative stability over environments seemed to favor polymorphisms in one or both of the competitors. A relatively less regular pattern of density or competitive response in series III was interpreted as evidence for the lack of coadaptedness between samples drawn from pure sites. Several limitations characteristic of such controlled studies were briefly discussed; however, the tentative conclusions from laboratory studies provided several promising clues for more critical field studies.This work was supported in part by a grant from the National Science Foundation (GB 8627).     

EFFECTS OF CANOPY POSITION AND IRRADIANCE ON THE LEAF PHYSIOLOGY AND MORPHOLOGY OF PENTACLETHRA MACROLOBA (MIMOSACEAE)

Pentaclethra macroloba (Willd.) Kuntze (Mimosaceae) is a dominant late-successional tree species in the Atlantic lowland forests of Costa Rica. Leaves of P. macroloba from three heights in the forest canopy were compared with leaves of seedlings grown in controlled environment chambers under four different irradiance levels. Changes in leaf characteristics along the canopy gradient paralleled changes resulting from the light gradient under controlled conditions. The effect of light or canopy position on light-saturated photosynthesis was small, with maximum photosynthesis increasing from 5 to 6.5 μmol m^−-2 s^−-1 from understory to canopy. Both chamber grown and field leaves showed large adjustments in photosynthetic efficiency at low light via reductions in dark respiration rates and increases in apparent quantum yields. Light saturation of all leaves occurred at or below 500 μmol m^−-2 s^−-1. Leaf thickness, specific leaf weight, and stomatal density increased to a greater extent than saturated photosynthesis with higher irradiance during growth or height in the canopy. As a result, there was a poor correspondence between leaf thickness and light-saturated photosynthesis on an area basis. It is concluded that Pentaclethra macroloba possesses the characteristics of a typical shade-tolerant species.     

Analysis of development of a subalpineAbies stand based on the growth processes of individual trees

The progress of growth of a subalpine youngAbies veitchii andA. mariesii forest during 25 years was analyzed on the basis of measurements of the processes of height growth of about 230 trees in a quadrat where the old canopy had been completely destroyed by a typhoon in 1959. The original forest floor sapling population had consisted of trees shorter than 2 m. Saplings grew faster after the breakage of the canopy than before,A. veitchii growing faster thanA. mariesii. During the 25 years of growth, a few well developed trees exceeded 6 m in height, while others remained around only 1 m or less. Some small trees, mostlyA. veitchii died at sites of high density. A bimodality in the distribution of tree height had developed with a trough at about 2.5 m, differentiating the trees into canopy and suppressed populations. Canopy trees grew with wide variation of rates, while most of the suppressed trees showed little recent growth. No difference was found in recent growth rates between the two canopyAbies species. Differences in height growth rates among individual canopy trees were analyzed on the basis of their horizontal crown overlapping. Competition models evaluating the difference in height between trees with overlapping crowns were shown to be effective. The height growth rate of a canopy tree appeared to be controlled by both the closely grown taller trees and the local density of trees including those shorter than the subject tree.     

Competition and patterns of resource use among seedlings of five tropical trees grown at ambient and elevated CO2

Summary Seedlings of five tropical trees, Cecropia obtusifolia, Myriocarpa longipes, Piper auritum, Senna multijuga and Trichospermum mexicanum, were grown both as individuals, and in competition with each other at ambient (350) and two levels of elevated CO₂ (525 and 700 l l^-1) for a period of 111 days. Growth, allocation, canopy architecture, mid-day leaf water potential and soil moisture content were assessed three times over this period for individually grown plants, and at the end of the experiment for competitively grown plants. In addition, leaf photosynthesis and conductance were assessed for the individually grown plants midway through the experiment, and light profile curves were determined for the competitive arrays at three stages of development. Elevated CO₂ did not affect photosynthesis or overall growth of the individually-grown plants but did affect canopy architecture; mean canopy height increased with CO₂ in Piper and Trichospermum and decreased in Senna. Stomatal conductance decreased slightly as CO₂ increased from 350 to 525 l l^-1 but this had no significant effect upon whole plant water use of leaf water potential. Soil moisture content for the individuals increased marginally as CO₂ increased, but this did not occur in the competitive arrays. There was a marked effect of CO₂ upon species composition of the competitive arrays; Senna decreased in importance as CO₂ increased while Cecropia, Trichospermum and Piper increased in importance. Stepwise regression analysis using competitive performance as the independent variable, and the various morphological and physiological parameters measured on the individually grown plants as independent variables, suggested that canopy height was the single most important variable determining competitive ability. Also significant were photosynthetic rate (particularly at low light levels) and allocation to roots early in the experiment. Light profiles in the canopy revealed that less than 15% of incident light penetrated to the level of mean canopy height. Results suggest that competition for light was the major factor determining community composition, and that CO₂ affected competitive outcome through its affect upon canopy architecture.This study was supported by a grant from the US Department of Energy     

Genetical analysis of chromosome 5A of wheat and its influence on important agronomic characters

Summary Chromosome 5A of bread wheat, Triticum aestivum carries the major gene, Vrnl, which is one of the main determinants of the winter/spring growth habit polymorphism in this species. Genetical analysis of this chromosome has been carried out using single-chromosome recombinant lines to establish the pleiotropic effects of this locus and two other major genes, q determining ear morphology and bl determining the presence of awns, on important agronomic characters. The three major genes were located on the long arm of chromosome 5A with a gene order of: centromere -bl-q-Vrnl. Analysis of quantitative characters from a winter sowing revealed pleiotropic effects of Vrnl or the effects of closely linked loci on the characters plant height, tiller number and spikelet number. However effects on ear emergence time were not associated with Vrnl but with q as were effects on spikelet number and ear length. In addition a locus determining yield/plant was located between Vrnl and q. Independant loci determining height and ear length were apparent on the short arm of chromosome 5A. From a spring sowing, however, there was a large pleiotropic effect of Vrnl on ear emergence time, as well as the effects previously detected. In addition, associated with q were effects on plant height and grain size which were not expressed from the winter sowing.     

Competition and sensitivity of wheat and wild oat exposed to enhanced UV-B radiation at different densities under field conditions

The influence of enhanced UV-B radiation (approximating a 15% ozone layer reduction) on competitive interaction between spring wheat (Triticum aestivum) and wild oat (Avena fatua) was examined in the field. The density-dependent mortality of both wheat and wild oat did not exhibit a significant difference between control and UV-B treatment conditions. A relatively high degree of competitive stress enhanced the effects of UV-B stress on biomass reduction. The relative competitive status of wheat in terms of total biomass increased under UV-B enhancement while it decreased when based upon grain production. Shifts in competitive balance occurred with significant changes in total biomass, especially when plants grew at higher densities in monocultures and mixtures. The sensitivity of wild oat to intensification of UV-B radiation at higher densities in mixtures was greater than that at lower densities. At all densities examined, wheat grown in mixture was significantly less sensitive to UV-B radiation than that in monoculture, and just the opposite for wild oat. The density of monocultures did not alter the response index (RI) of wheat and wild oat to enhanced UV-B radiation.     

Microplot studies with Xiphinema diversicaudatum under annual and perennial cropping regimes

Annual and perennial crops were grown in monoculture over three- or four-year experiments. Strawberry, raspberry and ryegrass maintained high population densities of Xiphinema diversicaudatum. Hop maintained relatively low population densities as did all annual crops tested, including winter wheat, winter beans, spring barley, potatoes, sugar beet and cabbage. Differences in nematode population density were not due to differences in soil porosity.     

Competitive interactions between the alien invasive annual grass <Emphasis Type="Italic">Avena fatua</Emphasis> and indigenous herbaceous plants in South African Renosterveld: the role of nitrogen enrichment

Nitrogen enrichment may play a role in successful invasion of indigenous South African mediterranean shrublands by alien invasive annual grasses. To test the hypothesis that an increase in nitrogen would result in a greater increase in biomass for an alien annual grass than for various indigenous plant functional groups, we conducted a field study in Western Cape Renosterveld shrubland fragments, surrounded by wheat or vinyards, to assess alien grass abundance in relation to soil nitrogen availability. Significant decreases in invasive annual grass Avena fatua cover and soil nitrogen were observed from the edges to the interior of Renosterveld habitat fragments and there was a significant positive relationship between Avena fatua cover and soil nitrogen. In addition, Avena fatua was grown in competition with three indigenous species of different functional types, an annual forb (Dimorphotheca pluvialis), a geophyte (Oxalis purpurea) and an indigenous perennial grass (Tribolium uniolae) at three concentrations of soil nitrogen in a pot experiment. Results revealed that the alien grass Avena fatua had significant increases in biomass when nitrogen was added, whereas the indigenous species did not. Alien grass competition significantly influenced performance of the annual forb and the indigenous grass but did not affect the geophyte. Results suggest the prioritization of hierarchical management options for the different functional groups in Renosterveld in response to invasive grasses. Nutrient enrichment through run off must be restricted to conserve remnant Renosterveld fragments.     

Earthworm communities in conventional wheat monocropping and low-input wheat-clover intercropping systems

A comparative study was conducted on earthworm communities in a conventional winter wheat monocropping system and a low‐input intercropping system in which successive crops of winter wheat were direct‐drilled into a permanent white clover sward. Earthworm abundance, biomass and species composition under the two cropping systems in the second and third years of successive cropping were assessed each spring and autumn in farm‐scale field plots at four sites using formalin and electrical extraction methods. The wheat‐clover cropping system supported larger earthworm communities (overall mean abundance 548 individuals m^?2, 137 g biomass m^?2) than conventional wheat monocropping (194 individuals m‐2, 36 g biomass m‐2). Between one and five more earthworm species were recorded in the wheat‐clover system than in the wheat system at three out of the four study sites. Wheat–clover cropping especially favoured species belonging to the epigeic and epigeic/anecic ecological groups such as Lumbricus castaneus, L. festivus, L. rubellus, juvenile Lumbricus and Satchellius mammalis. Earthworm communities in the wheat‐clover cropping system were comparable in size and species composition to communities normally found in perennial grassland‐type habitats such as pastures and grass‐legume leys.     

Comparative ecophysiology of two representativeQuercus species appearing in different stages of succession

Ecophysiological comparisons were made of the growth and photosynthetic characteristics between seedlings of deciduousQuercus serrata and evergreenQuercus myrsinaefolia. Q. myrsinaefolia seedlings naturally occurring in secondary coppice forests showed exponential-like growth in height with age, while sympatricQ. serrata seedlings were considerably smaller in height, their growth being limited by shortage of light. The photosynthetic characteristics measured under laboratory conditions showed no bases for the differences in growth between the two species on the forest floor: Light compensation points of the seedlings raised under 5% daylight were almost identical for the two species, being about 6.0 μE·m⁻²·s⁻¹. Growth analysis of seedlings planted in a coppice forest showed that bothQ. serrata andQ. myrsinaefolia could hardly grow during the summer under the shrub layer, when relative photon flux density (RPFD) was 0.9±0.5%. In the winter, when RPFD under the leafless canopy increased to 29.3±2.7%, the dry matter production of the evergreen seedlings ofQ. myrsinaefolia was much improved. Current-year seedlings of the species showed NAR of 0.102±0.021 g·dm⁻²·mo⁻¹ during the winter. Temperature dependency of photosynthesis and increment of leaf temperature by direct solar beam also indicated active photosynthesis ofQ. myrsinaefolia on the forest floor during the winter.     

Fungal communities and health status of roots of winter wheat cultivated after oats and oats mixed with other crops

Health status of winter wheat roots and thecomposition of wheat root fungi were studiedover 1996-1999 following the cultivation ofoats in a pure stand and mixed with otherplants as forecrops. The infection of wheatroots by >Gaeumannomyces graminis wasobserved to be largely dependent on the kind offorecrop; the best being oats in a pure stand,and then oats with pea or lupin mixtures. Inthe emergence and shooting phases, saprophyticfungi were dominant, while in the stage of harddough stage mainly pathogenic fungi, especially>G. graminis were common. The pathogenicfungi were mostly represented by >G.graminis and >Fusarium spp., while >Rhizoctonia spp. were much less frequent.The composition of the fungal communitydepended considerably on the forecrop anddevelopment phase of the plant. The kind offorecrop significantly affected the frequencyof infection by >G. graminis. The highestnumber of isolates was obtained from wheat rootsof crops grown after a mixture of oats andbarley.