Salinity tolerance of 'Valencia' orange trees on rootstocks with contrasting salt tolerance is not improved by moderate shade

The effects of shading in combination with salinity treatments were studied in citrus trees on two rootstocks with contrasting salt tolerance to determine if shading could reduce the negative effects of salinity stress. Well-nourished 2-year-old 'Valencia' orange trees grafted on Cleopatra mandarin (Cleo, relatively salt tolerant) or Carrizo citrange (Carr, relatively salt sensitive), were grown either under a 50% shade cloth or left unshaded in full sunlight. Half the trees received no salinity treatment and half were salinized with 50 mM Cl- during two 9 week salinity periods in the spring and autumn interrupted by an 11 week rainy period. The shade treatment reduced midday leaf temperature and leaf-to-air vapour pressure deficit regardless of salinity treatments. In non-salinized trees, shade increased midday CO2 assimilation rate (A(CO2)) and stomatal conductance, but had no effect on leaf transpiration (E(lf)). Shade also increased leaf chlorophyll and photosynthetic water use efficiency (A(CO2)/E(lf)) in leaves on both rootstocks and increased total plant dry weight in Cleo. The salinity treatment reduced leaf growth and leaf gas exchange parameters. Shade decreased Cl- concentrations in leaves of salinized Carr trees, but had no effect on leaf or root Cl- of trees on Cleo. There were no significant differences in leaf gas exchange parameters of shaded and unshaded salinized plants but the growth reduction from salinity stress was actually greater for shaded than for unshaded trees. Shaded trees on both rootstocks had higher leaf Na+ than unshaded trees after the first salinity period, and this shade-induced elevated leaf Na+ persisted after the second salinity period in trees on Carr. Thus, shading did not alleviate the negative effects of salinity on growth and Na+ accumulation.     

The Growth and Survival of Severely-shaded Tillers in Lolium perenne L.

The effect of shading a single tiller to below its compensationpoint for a period of 5 weeks in vegetative plants of Loliumperenne L. cv. S23, was studied in two different experimentseach employing two light regimes, one of which was common toboth experiments. In the first experiment tillers in the axils of the first leafwere shaded three weeks from appearance at both 40 and 70 Wm^–2. None of the shaded tillers died and they continuedto produce new leaves and increase in dry weight but at a reducedrate. In the second experiment, tillers with one emerged leafin any leaf axil position were shaded at 70 W m^–2 andin a treatment in which light was reduced to 13 W m^–2after initial growth at 70 W m^–2. As in the first experimentall shaded tillers survived at 70 W m^–2 but in the 70 13 W m^–2 transfer regime all shaded tillers died. In the second experiment shaded tillers in both light regimeswere supplied with ¹⁴C-assimilate by translocation from theremainder of the plant but in the 70 13 W m^–2 the initialsupport was withdrawn within 5 weeks of shading. The results are discussed in terms of the physiological relationshipsbetween the tillers of the grass plant. Lolium perenne L., growth of tillers, survival of tillers, effect of light     

Effects of Shading the First Leaf of Barley Plants on Growth and Carbon Nutrition of the Stem Apex

Effects of shading the first leaf on development of the apicalregion were investigated by examining the growth of leaf primordiaand the apical dome in the early seedling stages. Shade treatment affects the size of the dome; it was shown thatvalues for height, width, and volume of the dome of 12-day controlplants were always higher than for shaded plants. Primordialgrowth, in terms of length and dry weight, was reduced by shadeand growth in dry weight of the second, third, and fourth leaveswas shown to be dependent on photosynthetic production by thefirst leaf. Incorporation of ¹⁴C in the apical region was detected by autoradiographyon day 6 and increased with age. Transfer of assimilated carbonfrom the first leaf to the apex occurred during the first 3h after exposure to ¹⁴CO₂. On a unit dry-weight basis it isshown that the third and fourth leaves and apex incorporatedproportionately more labelled carbon than the larger older organssuch as the, second leaf. Shade treatment reduced incorporationinto the apical region and this is associated with the failureof the apex to grow over the period up to day 15. Evidence isprovided to show that in control plants the second leaf suppliescarbon to the apex from about day 12. The crucial importanceof the contribution of the first leaf to plant development isdiscussed.     

弱光胁迫对不同基因型玉米生长发育和产量的影响

以不同基因型玉米为材料,在玉米生长发育的3个主要阶段(苗期、穗期、粒期)进行分期遮光试验,研究不同时期弱光胁迫对不同基因型玉米生长发育和产量的影响。结果表明,遮光延缓了玉米叶片的出生速度,使叶片变薄;遮光可以延缓叶片的衰老,但遮光解除后则加速叶片的衰老;遮光造成植株高度增加,但恢复正常光照后,其株高却逐渐低于对照;遮光使干物质积累下降,抽雄吐丝日期推迟,尤其是吐丝日期推迟更多,并使产量降低,但不同基因型玉米不同遮光处理下降程度不同。试验的4个品种中,掖单2 2和豫玉2号受遮光影响较小,而掖单36 38和丹玉13受影响较大,即不同基因型玉米对弱光胁迫的敏感性不同     

Nitrogen reallocation and photosynthetic acclimation in response to partial shading in soybean plants

The first trifoliate of soybean was shaded when fully expanded, while the plant remained in high light; a situation representative for plants growing in a closed crop. Leaf mass and respiration rate per unit area declined sharply in the first few days upon shading and remained rather constant during the further 12 days of the shading treatment. Leaf nitrogen per unit area decreased gradually until the leaves were shed. Leaf senescence was enhanced by the shading treatment in contrast to control plants growing in low light. Shaded leaves on plants grown at low nutrient availability senesced earlier than shaded leaves on plants grown at high nutrient availability. The light saturated rate of photosynthesis decreased also gradually during the shading treatment, but somewhat faster than leaf N, whereas chlorophyll contents declined somewhat slower than leaf N.
Partitioning of N in the leaf over main photosynthetic functions was estimated from parameters derived from the response of photosynthesis to CO₂. It appeared that the N exported from the leaf was more at the expense of compounds that make up photosynthetic capacity than of those involved in photon absorption, resulting in a change in partitioning of N within the photosynthetic apparatus. Photosynthetic nitrogen use efficiency increased during the shading treatment, which was for the largest part due to the decrease in leaf N content, to some extent to the decrease in respiration rate and only for a small part to change in partitioning of N within the photosynthetic apparatus.     

Effects of decreasing source/sink ratio in soybeans on photosynthesis, photorespiration, transpiration and yield

Abstract. Shading of all side leaflets of a determinate soybean cultivar during pod filling significantly increased rates of photosynthesis in the unshaded centre leaflets, compared to centre leaflets of controls. Higher rates were associated with both higher stomatal and mesophyll conductances, and were reversible within 2 days when shades were removed. These higher rates of photosynthesis were not associated with decreased percentage enhancement by low oxygen, indicating that treatment effects were probably not associated with changes in photorespiration relative to photosynthesis. Percentage enhancement did, however increase as the plants approached physiological maturity, chiefly because of a decrease in photosynthesis.
In spite of these increases in rates of photosynthesis seed weight per plant was decreased by 37% in plants with side leaflets shaded for the entire pod-filling period and by 28% in plants shaded for only the second half of the period. In plants where shades were removed during the second half of pod filling seed yield was reduced by only 19% because shade removal delayed leaf senescence. The four treatments reduced yield by different mechanisms. Plants shaded continuously during pod filling produced fewer seeds than controls, but the weight per seed was similar. When shading was applied during the second half of pod fillings seed number was unchanged but weight per seed was significantly reduced. In contrast when shades were removed for the second half of pod filling, seed number remained similar to that of continuously shaded plants, but seed weight increased.
Although all shading treatments reduced yield, the reduction was not proportional to the 63% reduction in leaf area available for photosynthesis. This was because (1) photosynthetic rates in the centre leaflet of shaded plants were higher than rates in controls, (2) stem and lower surface photosynthesis in shaded leaf-lets contributed to whole leaflet photosynthesis.     

The photosynthesis of ryegrass leaves grown in a simulated sward

Plants were taken from simulated swards of perennial ryegrass (Lolium perenne) grown in a controlled environment and the rates of photosynthesis of the youngest fully expanded leaves, and the second and third youngest leaves on the same tillers were measured. The youngest leaves had the highest rates and the third the lowest, with the second leaves intermediate. The rate of photosynthesis in bright light of successive youngest expanded leaves decreased as the swards increased in leaf area, but did not when plants were grown so that the main stem was not shaded. When plants were grown at different densities and the photosynthetic rates of leaves of a particular ontogenetic rank were measured, it was found that leaves on plants from higher densities had lower rates of photosynthesis. Also leaves on plants grown in bright light had higher photosynthetic rates than those on plants grown in dim light. It is concluded that the decline in the photosynthetic capacity of successive leaves in a rapidly growing simulated sward is due to the intense shading to which they are subjected during their expansion.     

Effect of nitrogen deficiency on photosynthesis and the partitioning of 14C-labelled leaf assimilate in unshaded and partially shaded plants of Lolium temulentum

The effect of a deficiency of applied nitrogen on the rate of leaf photosynthesis, and on the subsequent partitioning of ¹⁴C-labelled leaf assimilate between new leaf, stem, tillers and root, was investigated in single plants of Lolium temulentum L., grown normally in controlled environments, or grown with collars shading the base of the plant. The nitrogen deficiency reduced the rate of leaf photosynthesis, increased the retention of assimilate in the leaf, suppressed the export of assimilate to tillers, and generally increased the export of assimilate to roots and to new leaves. Shading the base of the plant generally had little effect on the rate of leaf photosynthesis, reduced the export of assimilate to roots, and increased the export of assimilate to new leaf and to the stem, which elongated when shading was imposed.     

Effect of Defoliation on the Growth and Yield of Wheat

The effect of removal of the fifth (L₁), fifth and sixth (L₂) and fifth, sixth and seventh leaves (L₃) from the main shoot and of similar number of leaves from the primary and secondary tillers, on the growth and yield of two varieties of wheat, was investigated. The plants were grown in pots under natural conditions. Defoliation did not affect tillering, ear number, and number of spikelets per ear. The area of the last 3 leaves was reduced only in L₃. Defoliation reduced the growth of stem up to the flag leaf stage but thereafter there was a remarkable recovery in increase in height and dry weight. The grain yield of the main shoot was affected relatively more than that of the primary and secondary tillers by defoliation. The grain yield per plant was reduced by 5 per cent in L₁, and L₂ and by 11 per cent in L₃. The bearing of these observations on productivity per unit area is indicated and it is suggested that a plant with two fewer leaves per shoot may suffer less from mutual shading of leaves and hence prove more efficient in a dense community.     

不同生长阶段遮荫对番茄光合作用、干物质分配与叶N、P、K的影响

对番茄植株做了两种不同程度的遮荫处理,观测了夏季午间遮荫对光合速率,干物质积累量及其在根,茎,叶之间的分配,和叶N,P,K的含量以及经济产量的影响,发现不同时期遮荫影响不同。（1）遮荫增加三个阶段（开花早期,盛花期和开花后期）的气孔导度和胞间CO2浓度,显著降低开花早期中午的净化合速率,但盛花期中度遮荫（40%遮荫）使净光合速率随着时间的增加逐渐上升,在开花后期表现更加明显,平均净光合速率比对照高20%以上,蒸腾速率也增加较多。（2）开花早期和盛花期重度遮荫（如本实验中的75%遮荫）显著降低根,茎的干重,而开花后期中度遮荫的根,茎干重高于对照,但遮荫对叶干重的影响不明显。（3）开花早期和盛花期遮荫不明显影响叶片中N,P,K的含量,但开花后期中度遮荫使N,P,K含量增加,（4）开花早期两种遮荫对果实产量影响较小,但盛花期重度遮荫使产量降低,全部产量中无效部分所占的比例上升,开花后期中度遮荫的总产量和有效产量增加,单果重也增加,这些结果表明,在某些时期中度遮荫可以克服夏天辐射过强,气温过高对番茄的不良影响,对番茄生长,干物质积累和提高产量等有利,在生产上有意义。     

不同遮光处理对平茬后厚朴萌蘖株部分形态及生理指标的影响

以自然光照(遮光率0.0%)为对照,对遮光率20.0%、52.5%和78.6%条件下平茬后厚朴(Magnolia officinalis Rehd.et Wils)萌蘖株的部分形态和生理指标进行比较.结果表明:随遮光率提高,3个遮光处理组萌蘖株的株高呈现逐渐增大的趋势,萌蘖株叶中的叶绿素a、叶绿素b、类胡萝卜素和总叶绿素含量也基本上呈现逐渐增大的趋势,但萌蘖株的地径却呈现逐渐减小的趋势.萌蘖株的冠幅,单桩的萌蘖株数、叶片总数和叶芽总数,叶长和叶宽,单叶的干质量和叶面积,叶中叶绿素a含量与叶绿素b含量的比值(Chla/Chlb)和可溶性糖含量均在遮光率52.5%条件下最大,但仅冠幅、单桩叶片总数、单叶面积和叶中可溶性糖含量显著大于对照,分别较对照增加156.21%、28.18%、129.02%和64.31%;并且,多数指标在遮光率20.0%条件下居中、在遮光率78.6%条件下最小.然而,萌蘖株的叶长宽比、比叶质量及叶的相对含水量和可溶性蛋白质含量则在遮光率52.5%条件下最小,但仅比叶质量显著小于对照,较对照减少37.43%;并且,多数指标在遮光率20.0%条件下居中、在遮光率78.6%条件下最大.研究结果显示:适当的遮光处理能够促进平茬后厚朴萌蘖株的生长,遮光率52.5%为最优的遮光处理条件.     

Irradiance heterogeneity within crown affects photosynthetic capacity and nitrogen distribution of leaves in Cedrela sinensis

Because light conditions in the forest understory are highly heterogeneous, photosynthetic acclimation to spatially variable irradiance within a crown is important for crown‐level carbon assimilation. The effect of variation in irradiance within the crown on leaf nitrogen content and photosynthetic rate was examined for pinnate compound leaves in saplings of Cedrela sinensis, a pioneer deciduous tree. Five shading treatments, in which 0, 25, 50, 75 and 100% of leaves were shaded, were established by artificial heavy shading using shade screen umbrellas with 25% transmittance. Although the nitrogen content of leaves was constant regardless of shading treatment, ribulose 1·5‐bisphosphate carboxylase/oxygenase (Rubisco) content and light‐saturated photosynthetic capacity were lower in shade leaves within partially shaded crowns than within fully shaded crowns. Shade leaves within partially shaded crowns contained higher amount of amino acids. Most shade leaves died in partially shaded crowns, whereas more than half of shade leaves survived in totally shaded crowns. Assumptions on photosynthetic acclimation to local light conditions cannot explain why shade leaves have different photosynthetic capacities and survival rates in between partially and totally shaded crowns. Irradiance heterogeneity within the crown causes a distinct variation in photosynthetic activity between sun and shaded leaves within the crown.     

Tillering in grain sorghum over a wiide range of population densities: modelling dynamics of tiller fertility

The prediction of tillering is poor or absent in existing sorghum crop models even though fertile tillers contribute significantly to grain yield. The objective of this study was to identify general quantitative relationships underpinning tiller dynamics of sorghum for a broad range of assimilate availabilities. Emergence, phenology, leaf area development and fertility of individual main culms and tillers were quantified weekly in plants grown at one of four plant densities ranging from two to 16 plants m(-2). On any given day, a tiller was considered potentially fertile (a posteriori) if its number of leaves continued to increase thereafter. The dynamics of potentially fertile tiller number per plant varied greatly with plant density, but could generally be described by three determinants, stable across plant densities: tiller emergence rate aligned with leaf ligule appearance rate; cessation of tiller emergence occurred at a stable leaf area index; and rate of decrease in potentially fertile tillers was linearly related to the ratio of realized to potential leaf area growth. Realized leaf area growth is the measured increase in leaf area, whereas potential leaf area growth is the estimated increase in leaf area if all potentially fertile tillers were to continue to develop. Procedures to predict this ratio, by estimating realized leaf area per plant from intercepted radiation and potential leaf area per plant from the number and type of developing axes, are presented. While it is suitable for modelling tiller dynamics in grain sorghum, this general framework needs to be validated by testing it in different environments and for other cultivars.     

Improved growth and water use efficiency of cherry saplings under reduced light intensity

Cherry (Prunus avium L.) saplings were grown under natural sunlight (controls) or moderate shading (up to 30%, depending on the incident light intensity and the hour of the day). Reduced light intensity increased the dry mass of each of the plant components studied. Consequently, the total dry mass of shaded plants was significantly greater than that of controls at the end of the growing season. However, the diurnal trend in the level of photosynthesis (per unit of leaf area) of shaded plants was similar to the controls in August, but lower in September. As the growing season proceeded, reduced photosynthetic rates, thinner mesophyll and larger specific leaf area in the shaded plants indicated that leaf development had adapted to shaded conditions throughout the growing season. It is suggested that increased growth of shaded plants was caused by a higher initial relative growth rate and a greater whole-plant photosynthesis. Shading consistently reduced transpiration over the season, therefore improving water use efficiency of shaded leaves. Our results suggest that a moderate reduction in light intensity can be a useful method for improving growth and saving water in hot and dry environments.     

Functional significance of shade-induced leaf senescence in dense canopies: an experimental test using transgenic tobacco

Canopy photosynthesis models have predicted an optimal leaf area index (LAI; leaf area per unit surface area) and leaf nitrogen distribution at which whole-plant carbon gain per unit N is maximized. In this study we experimentally tested these models, using transgenic P(SAG12)-IPT tobacco (SAG; Nicotiana tabacum L.) plants with delayed leaf senescence and therefore a greater LAI and more uniform N distribution than the wild type (WT). In a competition experiment, the increased density of surrounding WT plants caused a greater reduction in dry mass of mature SAG target plants than in that of WT target plants, indicating negative effects of delayed leaf senescence on performance at high canopy density. Vegetative SAG plants achieved a lower calculated daily carbon gain than competing WT plants because the former retained leaves with a negative carbon gain in the shaded, lower part of the canopy. Sensitivity analyses showed that the carbon gain of SAG plants would increase if these lower leaves were shed and the N reallocated from these leaves were used to form additional leaf area at the canopy top. This strategy, which is adopted by the WT, is most advantageous because it results in the shading of competing neighbors.     

Response of soybean photosynthesis and chloroplast membrane function to canopy development and mutual shading

The effect of natural shading on photosynthetic capacity and chloroplast thylakoid membrane function was examined in soybean (Glycine max. cv Young) under field conditions using a randomized complete block design. Seedlings were thinned to 15 plants per square meter at 20 days after planting. Leaves destined to function in the shaded regions of the canopy were tagged during early expansion at 40 days after planting. To investigate the response of shaded leaves to an increase in available light, plants were removed from certain plots at 29 or 37 days after tagging to reduce the population from 15 to three plants per square meter and alter the irradiance and spectral quality of light. During the transition from a sun to a shade environment, maximum photosynthesis and chloroplast electron transport of control leaves decreased by two- to threefold over a period of 40 days followed by rapid senescence and abscission. Senescence and abscission of tagged leaves were delayed by more than 4 weeks in plots where plant populations were reduced to three plants per square meter. Maximum photosynthesis and chloroplast electron transport activity were stabilized or elevated in response to increased light when plant populations were reduced from 15 to three plants per square meter. Several chloroplast thylakoid membrane components were affected by light environment. Cytochrome f and coupling factor protein decreased by 40% and 80%, respectively, as control leaves became shaded and then increased when shaded leaves acclimated to high light. The concentrations of photosystem I (PSI) and photosystem II (PSII) reaction centers were not affected by light environment or leaf age in field grown plants, resulting in a constant PSII/PSI ratio of 1.6 ± 0.3. Analysis of the chlorophyll-protein composition revealed a shift in chlorophyll from PSI to PSII as leaves became shaded and a reversal of this process when shaded leaves were provided with increased light. These results were in contrast to those of soybeans grown in a growth chamber where the PSII/PSI ratio as well as cytochrome f and coupling factor protein levels were dependent on growth irradiance. To summarize, light environment regulated both the photosynthetic characteristics and the timing of senescence in soybean leaves grown under field conditions.     

环境因子对香根草生长习性的影响

环境因子对香根草生长习性的影响夏汉平,敖惠修,何道泉（中国科学院华南植物研究所,广州５１０６５０）ＥｆｆｅｃｔｏｆＥｎｖｉｒｏｎｍｅｎｔａｌＦａｃｔｏｒｓｏｎＶｅｔｉｖｅｒＧｒａｓｓＧｒｏｗｔｈ．￥ＸｉａＨａｎｐｉｎｇ;ＡｏＨｕｉｘｉｕ;ＨｅＤａｏｑ...     

Effects on the Stem of Winter Barley of Manipulating the Source and Sink during Grain-filling: I. CHANGES IN ACCUMULATION AND LOSS OF MASS FROM INTERNODES

Source and sink activities of field-grown barley plants weremanipulated by shading the whole plant or by de-earing at varioustimes after anthesis. The changes in the masses of the internodes,ears, grains and leaves were followed until the grain matured. Shading reduced the mass of ears and grains and, in addition,shading to 45% of incident radiation from anthesis preventedmature internodes from increasing in mass. Severe shading (to9% of incident radiation) from 11 d after anthesis caused thenormal decrease in mass of internodes to occur earlier thanin unshaded plants. Thus, only assimilate in excess of thatrequired for tructural growth was stored in internodes. Lowerinternodes of plants shaded from anthesis lost more of the massthat had been accumulated prior to anthesis than control plants,giving a greater potential pre-anthesis contribution to grainyield. Internodes from plants that were de-eared accumulated more massthan internodes of control plants in one year but not in another,i.e. the potential for internodes to store assimilate appearsto be exploited in some years but not others. The mass of internodesfrom de-eared plants either decreased after that of controlplants or their mass stayed constant. De-eared plants had anincreased number of tillers compared with control plants; thesetillers could have been an alternative sink for assimilate storedin the internodes of de-eared plants. Mobilization of assimilate from internodes is in response todemand for assimilate from grains being in excess of currentphotosynthesis. Key words: Winter barley, stem reserves, mobilization, shading, de-earing     

Effects of host shading on consumption and growth of the geometrid Epirrita autumnata: interactive roles of water, primary and secondary compounds

Shading is assumed to reduce allocation to plant phenolics and to defense in general. We here report the results of experimental shading of individual branches or whole canopies in mountain birch on foliar chemistry and on the growth and consumption of a geometrid, Epirritaautumnata. Branch‐wide shading tended to have at least as strong effects on both leaf chemistry and herbivore performance as canopy‐wide shading, indicating local responses of the host to shading. Responses to shading varied among the key leaf traits. Leaf water content was higher and toughness lower in shaded than in non‐shaded leaves. Leaf sugars were lower and protein‐bound and free amino acids higher in shaded than in control leaves. Sucrose and galactose were at high levels in unshaded branches adjacent to shaded ones, suggesting that partial shading enhanced translocation of sugars within canopies. Total phenolics and soluble proanthocyanidins were low in both shading treatments. Of the other phenolic groups, concentrations of gallotannins and cell‐wall‐bound proanthocyanidins did not differ between shaded and non‐shaded leaves. Epirrita larvae grew better in both types of shading treatments compared to either unshaded control trees or to unshaded branches in the branch‐shading trees. By far the most important correlate of larval growth was the amount of water consumed with leaf mass (r=0.94). When variance in water intake was standardized (also largely eliminating parallel variation in proteins), fructose and glucose still had significant positive correlations and proanthocyanidins negative with larval growth on control but not on shade leaves. Concentrations of several phenolic compounds correlated negatively with intake of dry matter and especially water, and different phenolics were important in shaded (gallotannins) and in control (flavonoids) leaves. Our findings strongly suggest that the effects of putatively defensive leaf traits on insect consumption and growth interact with nutritive leaf traits, particularly with water.     

The effect of shading on photosynthesis,growth, and regrowth following defoliation for Bromus tectorum

Summary The effect of full sunlight, 60%, or 90% attenuated light on photosynthetic rate, growth, leaf morphology, dry weight allocation patterns, phenology, and tolerance to clipping was examined in the glasshouse for steppe populations of the introduced grass, Bromus tectorum. The net photosynthetic response to light for plants grown in shade was comparable to responses for plants grown in full sunlight. Plants grown in full sunlight produced more biomass, tillers and leaves, and allocated a larger proportion of their total production to roots than plants grown in shade. The accumulation of root and shoot biomass over the first two months of seedling growth was primarily responsible for the larger size at harvest of plants grown in full sunlight. Plants grown under 60% and 90% shade flowered an average of 2 and 6 weeks later, respectively, than plants grown in full sunlight. Regrowth after clipping was greater for plants grown in full sunlight compared to those grown in shade. Even a one-time clipping delayed flowering and seed maturation; the older the individual when leaf area was removed, the greater the delay in its phenology. Repeated removal of leaf area was more frequently fatal for plants in shade than in full sunlight. For plants originally grown in full sunlight, regrowth in the dark was greater than for shaded plants and was more closely correlated to non-flowering tiller number than to plant size. This correlation suggests that etiolated regrowth is more likely regulated by the number of functional meristems than by differences in the size of carbohydrate pools. Thus, shading reduces the rate of growth, number of tillers, and ability to replace leaf area lost to herbivory for B. tectorum. These responses, in turn, intensify the effect of competition and defoliation for this grass in forests. B. tectorum is largely restricted to forest gaps at least in part because of its inability to acclimate photosynthetically, the influence of shade on resource allocation, and the role of herbivory in exacerbating these effects.