Shade avoidance alters the way plants grow, usually causing them to grow taller at the expense of placing resources into leaves, roots, seeds, and other harvestable materials. Sugar beet (Beta vulgaris L.) is a rosette-forming biennial species that has limited capacity to grow tall in the first year of growth. In the context of crop–weed competition, it is mostly unknown to what extent shade avoidance reduces yield in sugar beet relative to other effects like resource competition. To determine the extent of yield loss due to shade avoidance in a field-relevant situation, sugar beets were grown alongside Kentucky bluegrass (Poa pratensis L.) sod in a field study. Roots were separated with a steel root barrier placed into the ground between the grass and beets. Four treatments included a weed-free control (no root barrier or grass), a root barrier control (with root barrier but no grass), shade avoidance (with root barrier and grass), and full competition (with grass but no root barrier). The presence versus absence of grass was the primary driver of effects on measured sugar beet growth and yield parameters, regardless of whether a root barrier was present. Leaf number and root length were also impacted by the presence of the root barrier. These results suggest that shade avoidance is at least as important as root interactions and resource depletion in the context of early-season sugar beet yield loss due to weeds.

Scotch broom [Cytisus scoparius (L.) Link] is a large nonnative, leguminous shrub that threatens native plant communities by rapidly invading recently disturbed sites, competing vigorously for soil water and nutrients, and imparting soil legacy effects that inhibit native plants. In the Pacific Northwest, logging debris retention after forest harvesting prevents or slows C. scoparius invasions. A series of studies were conducted to determine potential mechanisms by which logging debris modifies the light environment to limit germination and growth of C. scoparius. In laboratory studies, seed germination did not vary significantly (P > 0.05): (1) between presence and absence of light for several temperature regimes, (2) when exposed to red (660-nm wavelength) versus far-red (730-nm wavelength) light, and (3) across a range of red/far-red light (R/FR) ratios. These results indicate that modification of the light environment by logging debris or plant canopies has little or no influence on C. scoparius germination. In a study to simulate effects of variable mass of logging debris, “heavy” debris (2 kg m−2) caused biologically relevant reductions in photosynthetic photon flux density (PPFD) and R/FR relative to conditions under “light” debris (1 kg m−2). Cytisus scoparius germination did not differ significantly between simulated heavy and light debris; however, values of seedling root and shoot biomass under heavy debris were 16% and 71% of those observed under light debris, respectively. These results indicate that heavy debris limits biomass of C. scoparius seedlings, particularly roots, by reducing both PPFD and R/FR, which increases seedling vulnerability to summer drought or other stressors. Retention of heavy logging debris after forest harvesting has potential application on sites likely to be invaded by C. scoparius, as well as those sites with seedbanks containing C. scoparius.

Field and greenhouse experiments were conducted from 2013 to 2015 at the University of Wyoming to evaluate the response of Beta vulgaris (L.) to reflected-light quality. Large-pail field studies included a factorial arrangement of three varieties of B. vulgaris (sugar beet, table beet, and Swiss chard) and reflected-light treatments (using either colored plastic mulch, grass, or bare-soil controls). Greenhouse studies included sugar beet as influenced by either grass or soil surroundings. In all studies, grass was grown in separate containers from B. vulgaris, so there was no root interaction. Grass was clipped regularly to prevent shading and competition for sunlight. Reflected light from different-colored plastic mulches (red, blue, green, black, clear) did not affect B. vulgaris growth. However, reflected light from the grass reduced the number of leaves in all B. vulgaris varieties such that there were 10 to 14 fewer leaves in B. vulgaris surrounded by grass compared with the soil treatment at 90 d after planting in the field study. Shade avoidance cues from surrounding grass reduced B. vulgaris total leaf area by 49% to 66%, leaf biomass by 21% to 30%, and root biomass by 70% to 72%. Similar results were observed in greenhouse experiments, where the grass treatment reduced sugar beet leaf biomass by 48% to 57% and root biomass by 35% to 64%. Shade avoidance cues have the potential to significantly reduce B. vulgaris yield, even in the absence of direct resource competition from weeds.

Laboratory studies were conducted to determine the effect of various factors on germination of dogfennel and yankeeweed. Dogfennel seed were found to be strongly photoblastic with no germination in the dark. Yankeeweed seed were moderately photoblastic, with 12% germination occurring in the absence of light Germination for both species increased in response to red light (650 nm), indicating phytochrome regulation. At the soil surface, dogfennel and yankeeweed emergence was 40 and 48%, respectively, but declined rapidly with increasing soil depths where dogfennel germination was lower than that of yankeeweed. Yankeeweed germination was 60 to 75% from 10 to 30 C while dogfennel germination was 45 to 70% between 15 to 30 C. Both species germinated over a broad range of pH (6 to 10) with the highest germination occurring at pH 8. Yankeeweed and dogfennel were moderately tolerant to water stress, but yankeeweed tolerated higher water stress than dogfennel. Both species germinated over a wide range of conditions, possibly enhancing adaptation to many diverse environments.

Light regulates many facets of plant growth and development through the effects of quantity of total energy and of photons, spectral quality, duration, and photoperiod. Numerous techniques and types of equipment are available for quantifying light in plant canopies. The effect of total quantity of light on weed and crop productivity has been described for many cropping systems. Recent work has focused on other aspects of light, in particular, spectral distribution of light (quality), transient light (sunflecks), and plant adaptation to changing light environments. The altered spectral quality of light in a plant canopy affects plant growth and morphology, which in turn affect competition for light. Dynamic plant response to transient light is also important to canopy photosynthesis and productivity. Plant physiological and morphological adaptation to fluctuating light is another potential factor regulating weed/crop interactions. Current cropping practices such as using smother crops and narrow row spacing exploit plant light responses to promote crop growth and suppress weed growth. A better understanding of plant responses to light quality, transient light, and fluctuating light environments will lead to a better understanding of how to manipulate the light environment in crop canopies to improve weed management.

Laboratory studies were conducted to determine the effect of various factors on germination of tropical soda apple. Tropical soda apple seed were found to be moderately photoblastic with 30% germination occurring in the dark. Germination increased in response to green (545 nm) and red light (650 nm) to 75 and 66%, respectively, indicating phytochrome regulation. No germination occurred in response to blue (450 nm) and far-red light (750 nm). Germination increased from 4 to 64% between 10 to 30 C, but no germination was found at 5 and 40 C. Maximum germination occurred at 30 C. A second degree polynomial (Y= −48.7 + 7.9X – 0.2X2, R2 = 0.76) best fit the data. Seed germinated over a broad range of pH (2 to 14) with the highest germination occurring at pH 8 and no germination occurring at pH 2 and 14 (Y = −63.3 + 30.5X – 1.9X2, R2 = 0.95). Seed germination declined as osmotic potential increased from 0 to −1.0 MPa (Y = 76.7 – 12.9X + 5.2X2, R2 = 0.87). Seedling emergence was maximum from depth of 3 to 6 cm, but no seedling emergence occurred when seed were planted below 12 cm (Y = 10.2 + 17.8X – 1.6X2, R2 = 0.93). Optimum germination occurred at a depth of 5.6 cm. Mechanical and sulfuric acid scarification increased the rate of germination but not the overall percentage. Tap water or hot water increased the rate of germination 26% and KNO3, GA3, or ethephon increased it 53%. Tropical soda apple seed germinated in response to variable environmental and edaphic conditions which could allow establishment in diverse ecosystems.

Germination of common mullein (Verbascum thapsus L.) in relation to constant and alternating temperatures, and light quality and duration was studied. In the dark, optimum germination occurred with 8-h warm periods of 30 to 40 C alternating with 16-h cold periods of 15 to 35 C. In the dark, no germination occurred below 10 C. With light during the 8-h warm periods, germination occurred at temperatures from 2 to 40 C, alternating with 0 C cold periods. The range of optimum temperatures for germination was also greatly expanded with light. Exposures to 24-h periods of red light, followed by incubation in the dark, produced the same germination response as 8 h of white light daily. Common mullein seeds germinated on the surface of the soil even though the seeds did not have mucilage.

Field studies were conducted in 2002 and 2003 to determine whether canopy formation influences pitted morningglory, common cocklebur, and sicklepod emergence in surface-tilled soybean. Each weed species was broadcast seeded before planting soybean in 19- and 97-cm-wide rows. Weed emergence beneath soybean was monitored after soybean emergence and compared with weed emergence in the absence of soybean (bareground treatment). Magnitude of daily diurnal soil temperature fluctuations diminished after soybean canopy formation, and light interception by soybean was positively related to the reduction in soil temperature. Canopy formation (50% light interception) occurred 16 to 17 d later in wide compared with narrow rows in both years. The red/far-red ratio of light available to seed on or near the soil surface was reduced from as much as 1.2 in full sunlight to less than 0.1 in the presence of a dense soybean canopy. Pitted morningglory emergence was not influenced by soybean canopy formation, whereas common cocklebur and sicklepod emergence were reduced as much as 33 and 68%, respectively. Although common cocklebur and sicklepod emergence diminished after soybean canopy formation, a small portion of the seedbank of both species emerged beneath the canopy. This research indicates that in a tilled system, emergence of some weed species is diminished by presence of an overlying canopy, but emergence does not completely cease with canopy formation. Late-season emergence of sicklepod and common cocklebur beneath a soybean canopy may contribute to replenishment of the soil seedbank, especially if these late-emerging cohorts are capable of surviving until the light environment is favorable for seed production.

Laboratory and greenhouse experiments were conducted to determine the effect of light, temperature, moisture stress, solution pH, and burial depth on coffee senna germination and emergence. Seeds germinated equally with or without light, and pretreatment with red or far-red light did not affect germination. Optimum temperature for germination was 25 C, and a high germination percentage (> 70%) occurred from 12.5 to 30 C. The low temperature threshold for germination was between 10 and 12.5 C, whereas the upper threshold was near 45 C. Coffee senna germination in response to moisture stress and solution pH differed at 15 and 30 C. At −0.4 MPa, no germination occurred at 15 C, whereas 15% germination occurred at 30 C. Optimum germination was at pH 6, but further increases in pH had a more negative effect on germination at 15 C than at 30 C. Coffee senna germination ranged from 9 to 12% at pH 3, but was 0% at pH 10, which indicates that coffee senna germination was more tolerant of acidic than basic solutions. Depth-mediated emergence inhibition was sigmoidal, with greatest emergence on the soil surface. Emergence from 2- to 10-cm depths reached 95% of the total emergence 1 to 3 d earlier in a sandy loam than in a sand soil. Mean emergence depth was 1.7 cm in the sand and 2.4 cm in the sandy loam soil. Knowledge gained from this research will be instrumental in developing a better understanding of the requirements for coffee senna germination and emergence, allowing further development and improvement of integrated weed management strategies specific to this troublesome weed.

A plant's ability to detect and adjust morphologically to changes in light quality (red–far-red [R:FR] ratio) is one mechanism by which a crop plant responds to weeds. To test this hypothesis, two experiments were conducted where corn was grown in growth cabinets under different light environments. First, to determine the effect of R:FR ratio on corn growth and development, treatments of high R:FR (1.37) and low R:FR (0.67) ratio were compared. These were established by planting corn in pots and then placing trays of either turface (a baked clay medium with high R:FR) or commercial grass sod (low R:FR) on each side of a row of corn pots. Grass sod was used to simulate low-growing weeds. The low R:FR sod treatment resulted in corn plants which were taller, had larger leaves, and greater shoot–root ratio than plants growing in the high R:FR turface treatment. In the second experiment, the effect of R:FR ratio on corn leaf azimuth position was examined. This was accomplished by adding a third treatment where each corn row had sod placed on one side and turface on the other. The proportion of leaves in four azimuthal classes was recorded. In the presence of sod, the proportion of leaves perpendicular to the corn row decreased, and this altered the proportion of leaves in other classes. Therefore, corn seedlings detected changes in light quality caused by the presence of sod (which simulated low-growing weeds) and responded by adjusting carbon allocation and leaf orientation to optimize the interception of light quantity and quality. These results support our hypothesis that low-lying vegetation can alter the growth of corn seedlings before competition for resources occurs. This change in growth may help explain the importance of early-season weed control in corn.

Characterizing the response of weeds to canopy shade is important for improved understanding of crop–weed competition and weed population dynamics. In 2000 and 2001, field studies were conducted in central New York state to examine the influence of three neighbor types (none, broccoli, or broccoli plus winter rye) and two locations (between or within rows of broccoli) on the morphology, phenology, and seed germination characteristics of Powell amaranth. Reductions in light availability and in the ratio of red-to-far red light were associated with increases in (1) partitioning of dry weight to stem tissue, (2) stem elongation, and (3) specific leaf area. Canopy shade also resulted in fewer main leaves at flowering and a reduced rate of leaf appearance but had no effect on the number of days to flowering. The relationship between Powell amaranth fecundity and aboveground dry weight was allometric, with both parameters declining significantly under competition. The weight of seeds produced did not vary significantly according to the competitive environment experienced by the maternal parent. However, the germination percentage of viable seeds was 40 to 50% lower for seeds maturing on plants grown under competition than without competition. Reductions in the number of main leaves at flowering and increased seed dormancy may be adaptive responses to canopy shade. Both mechanistic crop–weed competition models and population dynamic models would benefit from incorporation of data on the phenotypic plasticity of morphology, phenology, and seed germination characteristics of weeds.

Increased reflection of far-red light among plants can lower the red:far-red ratio (R:FR) of horizontally-propagated light before substantial shading among plants occurs. Although previous research suggests that altered R:FR can affect corn growth, it is not understood how early-season low R:FR (associated with higher plant densities in corn–weed communities than in weed-free corn) affects corn productivity in the field. We conducted field experiments from 2005 to 2007 to determine the effects of reduced R:FR on corn growth and yield. Corn was established at 53,800 plants ha−1 for a control R:FR treatment (weed-free corn) and at 107,600 plants ha−1 for a low R:FR treatment (simulated corn–weed community). The low R:FR treatment was thinned to the control plant density at V7 corn (seven fully-expanded leaf collars), which simulated total weed removal. Before thinning, R:FR (645:735 nm) at V7 corn in the low R:FR treatment (0.23) was less than 50% of that in the control R:FR treatment (0.49) across years. In 2005, stalk diameter, stem length, stem mass, leaf mass, specific leaf area, and total plant mass were less for V5 corn in the low R:FR treatment than in the control R:FR treatment. In 2006, V5 corn was taller in the low R:FR treatment than in the control R:FR treatment, but other morphological characteristics were similar between treatments. The shoot:root ratio of V5 (five fully-expanded leaf collars) (2005) and V7 corn (2006) was not affected by R:FR treatment, nor was shoot mass and grain yield of R6 corn (physiological maturity). Measurements in 2006 indicated that soil moisture and nutrient availability were not limiting and did not differ between treatments. Early-season R:FR in simulated corn–weed communities was similar to that in in situ corn–weed communities. These results suggest that low R:FR typical of early-season corn–weed communities was not associated with corn grain yield loss under field conditions in which water and nutrients were not limiting resources.

Pitted morningglory seed were collected in the fall of 2003 from Blackville, SC, and 2004 from Pendleton, SC, to assess the effect of After-Ripening and burial on light and temperature requirements for germination. Pitted morningglory germination was evaluated over a 12-mo period after maturation. Germination was neither stimulated by red light or inhibited by far-red light, nor was it reversible by red or far-red light. Light was not essential for germination of buried seed. Direct exposure to sunlight prevented germination of recently mature seed, but not once seed had sufficiently after-ripened. Pitted morningglory was capable of germination in darkness over a wide range of constant and fluctuating temperatures immediately after maturation. Germination in response to temperature varied with time of year after maturation, with the population from Pendleton having increased germination in May. Thermal fluctuations increased germination of both populations at suboptimal temperatures. Thermal amplitude regulation of germination varied over time and appeared to play a more important role in germination of after-ripened seed than recently mature ones. The ecological significance of changes in germination requirements with After-Ripening is discussed.

Experiments were conducted in 2002 and 2003 to determine the influence of soil thermal amplitude on common cocklebur emergence. Additionally, common cocklebur achenes were collected in fall of 2003 and of 2004 to assess changes in light and temperature requirements for germination over a 12-mo period under field conditions. Common cocklebur germination in response to the light environment and to constant and fluctuating temperatures were evaluated under controlled conditions following achene retrieval from the field. There was a linear inverse relationship between shade intensity and soil thermal amplitude, which explained 77% of the variability in emergence in the field over 2 yr. Emergence decreased as soil thermal amplitude declined, with 95% shading resulting in a 72 to 88% reduction in emergence. Neither red nor far-red light had much effect on germination, and burial did not induce a light requirement. Germination of achenes retrieved from the soil surface or buried in soil generally was not affected by red or far-red light, and the achenes did not acquire a red-light requirement following burial. Daily exposure to natural greenhouse light at 24 to 30 C was essential for germination immediately following achene maturation, whereas no germination occurred in darkness. A thermal fluctuation of 15 C increased germination percentages over those at constant temperatures regardless of time after maturation or retrieval-depth of achenes. The mean fluctuating temperature over all sample dates that was generally optimal for germination was 25 C (17.5/32.5 C low/high temperatures; 15 C daily fluctuation) in April or May and July or August in both years. Germination was generally optimum at constant temperatures of 35 or 40 C. The higher mean temperature requirement for germination at constant temperatures than at fluctuating temperatures likely contributes to reduced emergence in spring and summer months, when earlier emerging weed species or crops have already become established, and the thermal fluctuation requirement for germination reduces the likelihood of emergence in an environment where light availability is diminished.

Crop and weed competition studies rarely determine how plant-to-plant interactions alter the structure and physiology of crop roots. Soybean has the ability to detect neighboring weeds and to alter growth patterns including the allocation of resources to root growth. In this study, we hypothesized that low red : far red light ratio (R : FR) reflected from aboveground vegetative tissue of neighboring weeds would alter soybean root morphology and reduce root biomass and nodule number. All experiments were conducted under controlled conditions in which resources of light, water, and nutrients were nonlimiting. Low R : FR reflected from aboveground neighboring weeds reduced soybean seedling root length, surface area, and volume, including the number of nodules per plant. An accumulation of H2O2, an increase in malondialdehyde (MDA) content, a reduction in flavonoid content, and a decrease in 1,1-diphenyl-2-picrylhydrazyl (DPPH)–radicle scavenging activity were observed. The reduction in flavonoid content was accompanied by a decrease in the transcription of GmIFS and GmN93 and an increase in transcript levels of several antioxidant genes. These molecular and physiological changes may have a physiological cost to the soybean plant, which may limit the plant's ability to respond to subsequent abiotic and biotic stresses that will occur under field conditions.

Field experiments were conducted to determine whether exposure to reduced red : far-red light ratios (R : FR) typical of crop–weed environments was associated with adaptive changes in morphology, productivity, and fecundity of common lambsquarters. Plants were grown in reduced or ambient R : FR environments (both in full sunlight) until initiation of flowering, after which plants were grown in full sunlight or partial shade. At initiation of flowering, plants that had been exposed to reduced R : FR exhibited greater specific leaf area, stem elongation, main stem leaf area, specific stem length, and main stem mass compared with plants exposed to ambient R : FR. However, biomass allocation to stems, leaves, and roots did not differ between vegetative-stage R : FR treatments. At the end of flowering, morphology and productivity of plants exposed to partial shade did not differ between vegetative-stage R : FR treatments. In contrast, plants exposed to full sunlight during flowering after exposure to reduced R : FR during the vegetative stage had less total plant mass, less total leaf area, greater stem elongation, greater specific stem length, and a greater ratio of main stem to total stem mass compared with plants exposed to ambient R : FR during the vegetative stage. At physiological maturity, plants exposed to reduced R : FR during the vegetative stage and to partial shade during the reproductive stage had less total seed mass and fewer seeds compared with plants exposed to ambient R : FR during the vegetative stage and to partial shade during the reproductive stage. Fecundity of plants exposed to full sunlight during the reproductive stage did not differ between vegetative-stage R : FR treatments. These results indicate that exposure of common lambsquarters to reduced R : FR during the vegetative stage was maladaptive at later stages of growth in competitive environments, and suggest that interactions of light quality and quantity are important determinants of common lambsquarters fecundity.

Horseweed has become increasingly common and difficult to control in San Joaquin Valley vineyards, due in part, to the evolution of glyphosate resistance. The development of weed-suppressive vineyard designs in which the trellis design, spacing, and row orientation combine to cast dense shade on the weed canopy zone (WCZ) may reduce weed growth. The relevance of such a system to horseweed, which can grow to be as tall, or taller, than a typical grapevine trellis, is uncertain. Also unknown is whether a glyphosate-resistant (GR) biotype and glyphosate-susceptible (GS) biotype would perform similarly under such conditions. Therefore, we compared the growth and development of two potted horseweed biotypes (GR and GS) in vinerows oriented east–west (EW) and north–south (NS). Rows oriented EW allowed less light penetration to the WCZ than NS rows throughout the study, and horseweed biotypes responded to low light levels by producing leaves with larger specific leaf area and leaf area ratios than those in the NS rows. Also, the leaf, stem, and root dry weight of the horseweed plants in the EW rows was reduced by 30% compared to the horseweed plants in NS rows. Leaf number was also reduced in the horseweed plants in the EW rows, but only for the GS biotype. Row orientation did not affect phenological development or the number of seeds produced by the GR or GS biotypes, but the GR biotype budded, flowered, and set seed approximately 1 wk earlier than the GS biotype. Thus, shade associated with the EW vinerows reduced horseweed growth, but not fecundity, and the GR biotype reached reproductive maturity earlier than the GS biotype.

Previous studies have suggested that the reduction in the root/shoot ratio that accompanies the shade avoidance response may reduce the tolerance of individuals to subsequent nutrient or moisture limitations. In this work, we examined the impact of the shade avoidance response on maize seedling growth and development and the response of these plants to a subsequent abiotic stress. Seedlings were grown in a field fertigation system under two light quality environments, ambient and a low red to far-red ratio, which were designed to simulate weed-free and weedy conditions, respectively. This system also enabled the controlled restriction of water and nutrients, which reduced the relative growth rate of the crop and created a secondary stress. Results of this study indicate that, while the shade avoidance response did reduce the root/shoot ratio in maize, this effect did not reduce plant tolerance to subsequent abiotic stress. Rather, the apparent additivity or synergism of shade avoidance and the secondary stressor on yield loss depended on whether the net effect of these two stressors was sufficiently large to shift the population toward the point where reproductive allometry was broken.

Traditional shade house experiments that expose plants to relatively uniform irradiance and light quality are inadequate to characterize the morphological, allocational and physiological plasticity that seedlings show to different gap environments. Here the design of a pot experiment is described that simulates the daily time course of irradiance and light quality in idealized gaps of six different sizes. Differences in response to gap size are illustrated using data from two pioneer species, Ochroma pyramidale, which recruits exclusively in large gaps and clearings, and Luehea seemannii, which colonizes small branchfall gaps as well as large gaps. Ochroma outperformed Luehea in relative growth rate in all except the smallest simulated gap size. Ochroma's superior performance in the larger gaps could be attributed to a larger proportional investment in leaf biomass (i.e. a higher leaf area ratio, LAR), and higher photosynthetic rates both on a leaf area and leaf mass basis. In the smallest simulated gaps LAR was not significantly different between the species, but Ochroma maintained a higher net assimilation rate. These results fail to support the suggestion that gap partitioning among pioneer species arises directly from morphological and biochemical specialization to particular gap light environments. Instead, it is suggested that partitioning may result from a trade-off between seedling growth and mortality determined by species allocational patterns and mediated by interactions with herbivores and pathogens.

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[ratio ]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.