Physiological, morphological and allocational plasticity in understory deciduous trees: importance of plant size and light availability

In a 4-year study, we investigated changes in leaf physiology, crown morphology and whole-tree biomass allocation in seedlings and saplings of shade-tolerant sugar maple (Acer saccharum Marsh.) and intermediate shade-tolerant yellow birch (Betula alleghaniensis Britt.) growing in natural understory light (0.5 to 35% of full sunlight) or in understory light reduced by 50% with shade nets to simulate the effect of gap closure. Leaf physiological parameters were mainly influenced by the light gradient, whereas crown morphological and whole-tree allocational parameters were mainly influenced by tree size. No single physiological, morphological or allocational trait was identified that could explain the difference in shade tolerance between the species. Yellow birch had higher growth rates, biomass allocation to branches and leaf physiological plasticity and lower crown morphological plasticity in unmodified understory light than sugar maple. Sugar maple did not display significant physiological plasticity, but showed variation with tree size in both crown morphology and whole-tree biomass allocation. When sugar maple was small, a greater proportion of whole-tree biomass was allocated to roots. However, physiological differences between the species decreased with decreasing light and most morphological and allocational differences tended to disappear with increasing tree size, suggesting that many species differences in shade-tolerance are expressed mainly during the seedling stage. Understory trees of both species survived for 4 years under shade nets, possibly because of higher plasticity when small and the use of stored reserves when taller.     

Growth of Festuca pallescens in Silvopastoral Systems in Patagonia, Part 2: Parameterization of Models of Stomatal Conductance and Leaf Photosynthesis

A combined model of stomatal conductance and photosynthesis was developed for Festuca pallescens (St. Ives) Parodi, a forage species in Patagonia. Curves showing the relationship between photosynthesis and photosynthetic photon flux density (PPFD) were constructed for plants grown under differing levels of water availability, relative humidity (RH) and air temperature (T). Stomatal conductance (g_s) was related to these variables and pre-dawn leaf water potential (ψ_pd) using an empirical multiplicative submodel. Parameters of the photosynthesis-PPFD curves were related to the average g_s values for each curve to introduce stomatal limitation on photosynthesis. Considering the simplicity of the models, estimated stomatal conductance and photosynthesis agree satisfactorily with independent measured values in the field and in the glasshouse, particularly in the range of low and medium values of both variables (R² = 0.84 and 0.87 for g_s and photosynthesis models, respectively). Photosynthesis–PPFD curves were also determined under field conditions for plants growing under shade and in the open, in a silvopastoral trial in northwestern Patagonia. No significant differences in the photosynthetic light response curves were found between these locations, but slight increases in maximum assimilation rate and quantum yield (light use efficiency) were found for leaves grown under shade. This study of environmental influences on photosynthesis in F. pallescens may help to predict its capacity to grow under trees in silvopastoral systems. In addition, this simple model may be easily parameterised for other species to predict photosynthetic responses under different environmental conditions.     

Influence of light availability on growth, leaf morphology and plant architecture of beech ( Fagus sylvatica L.), maple ( Acer pseudoplatanus L.) and ash ( Fraxinus excelsior L.) saplings

In a field study, we measured saplings of beech, ash and maple growing in a fairly even-aged mixed-species thicket established by natural regeneration beneath a patchy shelterwood canopy with 3–60% of above canopy radiation reaching the saplings. Under low light conditions, maple and ash showed a slight lead in recent annual length increment compared with beech. With increasing light, ash and maple constantly gained superiority in length increment, whereas beech approached an asymptotic value above 35% light. A suite of architectural and leaf morphological attributes indicated a more pronounced ability of beech to adapt to shade than ash and maple. Beech displayed its leaves along the entire tree height (with a concentration in the middle crown), yielding a higher live crown ratio than ash and maple. It allocated biomass preferentially to radial growth which resulted in low height to diameter ratios, and expressed marked plagiotropic growth in shade indicating a horizontal light-foraging strategy. In addition, beech exhibited the highest specific leaf area, a greater total leaf area per unit tree height, a slightly greater leaf area index, and a greater plasticity to light in total leaf area. Ash and maple presented a “gap species” growth strategy, characterized by a marked and constant response in growth rates to increasing light and an inability to strongly reduce their growth rates in deep shade. In shade, they showed some plasticity in displaying most of their leaf area at the top of the crown to minimize self-shading and to enhance light interception. Through this, particularly, maple developed an “umbrella” like crown. These species-specific responses may be used for controlling the development of mixed-species regeneration in shelterwood systems.     

Crown architecture of understory and open-grown white pine (Pinus strobus L.) saplings

Crown architecture and growth allocation were studied in saplings of eastern white pine (Pinus strobus L.), a species classified as intermediate in shade tolerance. A comparison was made of 15 understory saplings and 15 open-grown saplings that were selected to have comparable heights (mean of 211 cm, range of 180-250 cm). Mean ages of understory and open-grown trees were 25 and 8 years, respectively. Understory trees had a lower degree of apical control, shorter crown length, and more horizontal branch angle, resulting in a broader crown shape than that of open-grown trees. Total leaf area was greater in open-grown saplings than in understory saplings, but the ratio of whole-crown silhouette (projected) leaf area to total leaf area was significantly greater in understory pine (0.154) than in open-grown pine (0.128), indicating that the crown and shoot structure of understory trees exposed a greater percentage of leaf area to direct overhead light. Current-year production of understory white pine was significantly less than that of open-grown white pine, but a higher percentage of current-year production was allocated to foliage in shoots of understory saplings. These modifications in crown structure and allocation between open-grown and understory white pine saplings are similar to those reported for more shade-tolerant fir (Abies) and spruce (Picea) species, but the modifications were generally smaller in white pine. As a result, white pine did not develop the flat-topped "umbrella" crown structure observed in understory fir and spruce, which approaches the idealized monolayer form that maximizes light interception. The overall change to a broader crown shape in understory white pine was qualitatively similar, but much more limited than the changes that occurred in fir and spruce. This may prevent white pine from persisting in understory shade as long as fir and spruce saplings.     

Hydraulic function contributes to the variation in shoot morphology within the crown in Quercus crispula

Hydraulic and light environments have variation within the crown in well-grown trees. Shoot morphology and shoot hydraulics were compared between the upper and lower crown or among branching patterns in well-grown Quercus crispula Blume. Shoots in the upper crown had longer and thicker axes and lower water potential than did shoots in the lower crown. Hydraulic conductance from the soil to the shoot did not differ between the upper crown and the lower crown. Shoots in the upper crown are exposed to hydraulic stress, and shoots in the lower crown are under shade stress. Shoot morphology and shoot hydraulic traits (i.e., higher Huber value and higher hydraulic conductivity) in the upper crown affected the hydraulic conductance of shoots. Shoots in the lower crown showed larger light-receiving leaf area per leaf biomass investment, which is an adaptive morphology under shaded environments. Shoot morphology and shoot hydraulics were not correlated to branching pattern significantly, but shoots with higher branching intensity in the upper crown represented trends for higher hydraulic conductivity. These results reveal that shoot morphological and physiological characteristics in the upper crown reduce hydraulic stress, and those in the lower crown reduce shade stress. I conclude that vertical position within a crown affects both morphological and physiological acclimation for light acquisition and hydraulic conductance, and that hydraulic architecture is associated with crown architecture.     

Growth of <Emphasis Type="Italic">Dactylis glomerata</Emphasis> along a light gradient in the central Appalachian region of the eastern USA: II. Mechanisms of leaf dry matter production

Microsite influences development and resource allocation of Dactylis glomerata L. (orchardgrass), a traditional pasture species with potential as an understory crop in silvopasture of humid temperate regions. An experiment using container-grown orchardgrass was conducted under field conditions to determine how open (O), shaded woodland (W) and open-to-shaded woodland transition zone (E _O, E _W) microsites influenced leaf DM production. Plants established in spring (SP) and late summer (LS) were clipped each time mean canopy height reached 20 cm. Dry matter production and allocation among structures differed, as a function of light attenuation. Specific leaf area (SLA) and photosynthetic nitrogen-use efficiency (PNUE) were associated with leaf DM production, whereas leaf N, net assimilation rate and shoot total nonstructural carbohydrates (TNC), were not. Specific leaf area was related to leaf DM of LS plants, whereas PNUE influenced leaf DM of SP plants. Stembase TNC was inversely related to relative regrowth rate (RG_RR) with RG_RR greatest and TNC the least at W. The relationship for RG_RR and TNC for SP plants growing at O and LS plants growing at W was similar. Regardless of how indices of growth are related, SP and LS plantings responded as separate populations (representing young and established plants respectively) that have different leaf DM production efficiencies. Orchardgrass was able to sustain leaf production when subjected to simultaneous stresses of shade and repeated defoliation. The LS plants growing at W respond in a manner similar to SP plants and may require management practices attuned to establishing or immature plants. The US Government’s right to retain a royalty-free, non-exclusive copyright is achnowledged.     

遮阴处理对金蒲桃生长量及光合参数的影响

以2 a生金蒲桃(Xanthostemon chrysanthus)幼苗为材料,研究了全光照,50%遮阴,70%遮阴,90%遮阴4种遮阴处理对金蒲桃生长特性、生物量分配、光合生理变化的影响,结果表明:与自然光全处理对比,遮光极显著(P<0.01)地抑制了金蒲桃幼苗生物量的积累,遮光处理组叶片数量减少,叶生物量比下降,茎生物量比增加;光补偿点、光饱和点、暗呼吸速率和最大光合速率降低,光能利用效率低于全光照叶,强光下遮阴叶的净光合速率保持稳定.上述结果说明:遮阴处理后,金蒲桃在生长特性、生物量分配和光合参数上表现出对遮阴弱光的适应,表明金蒲桃是一种能广泛应用于各种光照条件的优良景观树种.     

不同栽植密度下欧美杨叶片耐荫性与生物累积量的关系

为阐明不同栽植密度下欧美杨无性系树冠叶片耐荫性与生物累积量的关系,选择高密度(2 m×3 m)和低密度(2 m×5 m)栽植的不同亲本的5个欧美杨无性系为研究对象,测量了生物量、树冠结构、叶面积指数及光合参数。结果表明:(1)不同无性系对栽植密度有不同的反应;(2)具有较高生物量的欧美杨无性系树冠叶片在高、低栽植密度下均表现出较强的耐荫性特征;(3)树冠耐荫性在不同密度中表现出差异性,在低密度栽植中,树冠上层耐荫性与生物量和叶面积指数关系密切;在高密度栽植中,树冠上、下层叶片耐荫性与生物量和叶面积指数关系密切;(4)树冠耐荫性特征受亲本和环境的共同影响,由低密度到高密度,具有较高生物量的03-04-97、03-04-111、03-04-171树冠中下层叶片光补偿点和暗呼吸速率大幅度降低。该研究结果可为选择适合高密度栽植的欧美杨无性系提供数据和理论支持。     

Morphological plasticity and regeneration strategies of velvet leaf blueberry (Vaccinium myrtilloides Michx.) following canopy disturbance in boreal mixedwood forests

The effects of canopy disturbance on the abundance, growth, morphological plasticity, biomass allocation and fruit production of velvet leaf blueberry (Vaccinium myrtilloides Michx.) were examined in 1996 in a second-growth boreal mixedwood forest near Nipigon, northwestern Ontario that had been logged by either shelterwood cutting or clearcutting in 1993. We found that V. myrtilloides was able to persist in both open and closed canopy boreal mixedwood forests managed for commercial timber extraction. Persistence under heavy shade conditions was accompanied by significant morphological and biomass allocation plasticity. Specific leaf area, leaf area, individual leaf weight, and the proportion of total biomass in stems and foliage changed along an understory light gradient from 0% to 67% percent photosynthetic photon flux density (% PPFD). The degree of above-ground morphological plasticity may explain blueberry's ability to survive under low light conditions. Reproductive performance of V. myrtilloides was greatest under the partial shade conditions associated with shelterwood cutting. Blueberry bushes growing in clearcuts overgrown with 3-year old aspen (Populous tremuloides Michx.) saplings remained mostly vegetative whereas the number, fresh weight and dry weight of berries in shelterwood cuts was 94% grater than that produced after clearcutting. We attributed the lower fruit yields in the clearcuts to heavy shading from regenerating hardwoods, and mechanical damage to above-ground biomass. The paucity of seedling regeneration as well as extensive mechanical damage to above-ground stems by logging equipment delayed vegetative regeneration of V. myrtilloides in large canopy openings of the clearcut blocks. Unlike other more aggressive ericaceous species (e.g. Kalmia angustifolia var. angustifolia L., Gaultheria shallon Pursh.), V. myrtilloides was unable to resist invasion from faster growing hardwood species (e.g. P. tremuloides) and was rapidly overtopped. V. myrtilloides plants in the uncut control blocks received 3.9% of full sunlight, whereas those growing in the partial cut and clearcut blocks received an average of 25.3% and 32.5% PPFD, respectively. Cover of vegetation over-topping blueberry plants was highest in the uncut forest (90.3%), but was not significantly different between the partial cut (45.5%) and clearcut (50.1%) treatment blocks.     

Effect of shade on biomass accumulation and partitioning for Eucalyptus camaldulensis sprouts

ABSTRACT

Eucalyptus is widely cultivated in homogeneous monocultures throughout the world alongside plants of the same age. Furthermore, the productivity of forest stands is highly dependent on how solar radiation is intercepted and used. Meanwhile, it is possible to consider cultivating trees of different ages in the same area, and thus under different regimes of available solar radiation, creating a sequence of plantings and harvestings and to develop Eucalyptus-based agroforestry systems. We have examined biomass accumulation of Eucalyptus camaldulensis along a shade gradient. Eucalyptus trees presented power-law responses to increasing irradiance for biomass, indicating it has low plasticity and is unable to maintain growth under lower irradiance levels. Biomass partitioning showed no dependence on available irradiation. Leaf density presented very little increase toward sunnier places meaning that an increment in Eucalyptus biomass would be mostly due to its increase in size and thus higher radiation interception. Careful management of shade will be a key consideration for the integration of Eucalyptus into agroforestry. With knowledge of the growth habits of different species, we will have the potential to propose more perennial ways in which to manage land and reduce anthropogenic disturbances by avoiding clear-cuts of areas that removes all plant structures.     

The impact of shade on morphology,growth and biomass allocation in <Emphasis Type="Italic">Picea sitchensis</Emphasis>, <Emphasis Type="Italic">Larix</Emphasis> × <Emphasis Type="Italic">eurolepis</Emphasis> and <Emphasis Type="Italic">Thuja plicata</Emphasis>

To determine the effect of shade on morphology, growth and biomass allocation in Picea sitchensis, Larix × eurolepis and Thuja plicata, seedlings were grown in the open or under shadehouses providing 25%, 50% and 75% reductions of full-light for two growing seasons. For most of the characteristics assessed there was no significant interaction between species and shade indicating that the morphological responses to changing shade treatments were not species-dependent. After two growing seasons the mean height increment for the three species was significantly greater in 25% (76.1 cm) and 50% shade (74.9 cm) than in the open (69.5 cm). Root collar diameter increment, shoot, root and total biomass declined significantly with increasing shade while the opposite was true for the height:diameter ratio. In both western red cedar and hybrid larch the shoot:root ratio was significantly greater in the shade while in Sitka spruce this characteristic was not influenced by shade. While all species had significantly greater specific shoot areas in 75% shade than in 0% shade, this trend was particularly pronounced in hybrid larch. In hybrid larch and western red cedar, the normalised specific projected shoot area increased significantly with increasing shade. The opposite trend was observed for Sitka spruce. We conclude that in the main the species studied demonstrated similar shade acclimation responses despite their reported differences in shade tolerance.     

Differences in biomass allocation patterns between saplings of two co-occurring Mediterranean oaks as reflecting different strategies in the use of light and water

Production and biomass allocation patterns, the growth rates of aboveground biomass, and crown traits were examined in saplings of the deciduous Quercus faginea and the evergreen Q. ilex to determine whether differences in these traits might account for the greater mortality during periods of drought undergone by Q. faginea. Strong differences were observed in almost all the traits analyzed, which suggests that the two species use different strategies to cope with the main limiting factors for woody seedling establishment in Mediterranean environments: excess light and low water availability. In Q. faginea, sapling design seems to be oriented to maximize light capture and, hence, leaf productivity during the short life span of the leaf biomass. Thus, the seedlings of Q. faginea showed crown traits that permit self-shading to be minimized: longer shoots with more spaced leaves that result in lower leaf area index than in Q. ilex. In addition, the larger area per unit leaf biomass in Q. faginea leads to a larger interceptive leaf area per unit plant mass and to higher light capture. These characteristics imply higher investments in woody tissues (SWR) that permit the plants to support a wide canopy and facilitate water transport to meet the strong transpiratory demands of a canopy with such characteristics. By contrast, in Q. ilex, saplings are apparently designed to guarantee leaf survival against temperature extremes and photoinhibition through avoidance of excessive radiation.     

Growth and photosynthetic responses of four Virginia Piedmont tree species to shade

To determine the effects of shade on biomass, carbon allocation patterns and photosynthetic response, seedlings of loblolly pine (Pinus taeda L.), white pine (Pinus strobus L.), red maple (Acer rubrum L.), and yellow-poplar (Liriodendron tulipifera L.) were grown without shade or in shade treatments providing a 79 or 89% reduction of full sunlight for two growing seasons. The shade treatments resulted in less total biomass for all species, with loblolly pine showing the greatest shade-induced growth reduction. Yellow-poplar was the only species to show increased stem height growth in the 89% shade treatment. The shade treatments increased specific leaf area of all species. Quantum efficiency, dark respiration and light compensation point were generally not affected by the shade treatments. Quantum efficiency, dark respiration, maximum photosynthesis and light compensation point did not change consistently between the first and second growing seasons. We conclude that differences in shade tolerance among these species are not the result of changes in the photosynthetic mechanism in response to shade.     

Shoot structure and growth along a vertical profile within a Populus-Tilia canopy

We investigated shoot growth patterns and their relationship to the canopy radiation environment and the distribution of leaf photosynthetic production in a 27-m-tall stand of light-demanding Populus tremula L. and shade-tolerant Tilia cordata Mill. The species formed two distinct layers in the leaf canopy and showed different responses in branch architecture to the canopy light gradient. In P. tremula, shoot bifurcation decreased rapidly with decreasing light, and leaf display allowed capture of multidirectional light. In contrast, leaf display in T. cordata was limited to efficient interception of unidirectional light, and shoot growth and branching pattern facilitated relatively rapid expansion into potentially unoccupied space even in the low light of the lower canopy. At the canopy level, T. cordata had higher photosynthetic light-use efficiency than P. tremula, whereas P. tremula had higher nitrogen-use efficiency than T. cordata. However, at the individual leaf level, both species had similar efficiencies under comparable light conditions. Production of new leaf area in the canopy followed the pattern of photosynthetic production. However, the species differed substantially in extension growth and space-filling strategy. Light-demanding P. tremula expanded into new space with a few long shoots, with shoot length strongly dependent on photosynthetic photon flux density (PPFD). Production of new leaf area and extension growth were largely uncoupled in this species because short shoots, which do not contribute to extension growth, produced many new leaves. Thus, in P. tremula, the growth pattern was strongly directed toward the top of the canopy. In contrast, in shade-tolerant T. cordata, shoot growth was weakly related to PPFD and more was invested in long shoot growth on a leaf area basis compared with P. tremula. However, this extension growth was not directed and may serve as a passive means of avoiding self-shading. This study supports the hypothesis that, for a particular species, allocation patterns and crown architecture contribute as much to shade tolerance as leaf-level photosynthetic acclimation.     

紫茎泽兰(Eupatorium adenophorum)对光强的生态适应性

紫茎泽兰(Eupatorium adenophorum)是我国主要外来入侵植物之一,在我国西南地区迅速传播,引起了巨大的经济损失。本文比较研究了不同光强下生长14个月的(透光率RI分别为10%、20%、30%、55%、100%)紫茎泽兰幼苗的生物量分配、叶片形态和生长反应特性。结果表明:紫茎泽兰在不同的光强条件下显示出极强的叶生态适应性。植株总生物量随光强升高而增加(RI从10%到55%),但在全光照下总生物量反而有所下降。株高也随光强增大(RI从10%到30%)而增大,但到达一定程度(RI30%)后,株高反而下降。在弱光照条件下,从比叶面积(SLA),叶面积比(LAR)和比茎长(SSL)等指标表明,植物通过叶片变薄、变大增加单位生物量的叶面积来捕获光能。随着光强的升高,平均相对生长速率增加,在本试验尺度下,RI在55%左右达到平均相对生长速率最大值。植株在RI为30%–55%下生长状况好于全光照条件下的幼苗。在高遮荫条件下能阻止其它生物的生存和生长,这可能是紫茎泽兰强入侵性的一种生态适应策略。结论:紫荆泽兰对光照强度拥有极强的生态适应性,可能是紫荆泽兰入侵性比较强的重要原因之一。图3参26。     

Biomass allocation and assimilation efficiency of naturalTilia amurensis samplings in response to different light regimes

Biomass allocation and assimilation efficiency of natural Amour linden (Tilia amurensis) samplings in different light regimes were analyzed in the paper. The results showed that shoot increment of samplings in gap was the highest and that of samplings under canopy was the least. Samplings in gap expressed apical dominance strongly but samplings in full sun and under canopy behaved intensive branching. Lateral competition or moderate shading was favored to bole construction. The patters of biomass allocation of samplings in different light environment were rather similar. The biomass translocated to stem was more than that to other organs, and about one half of photosynthate was used to support leaf turn over. On the contrary, photosynthates of samplings in full sun were mostly consumed in leaves bearing and energy balancing. The carbon assimilation for leaves of samplings in gap was the most efficient, and more carbons were fixed and translocated to non-photosynthetic organs, especially to stemwood. Responsible editor: Zhu Hong     

Structural scaling of light interception efficiency in Picea engelmannii and Abies lasiocarpa

Sunlight interception efficiency was compared at the leaf, shoot, branch and crown levels for Picea engelmannii (Parry) and Abies lasiocarpa ((Hook.) Nutt.), dominant tree species of the central Rocky Mountains, USA. The ratio of silhouette to total leaf area (STAR) was used to quantify the efficiency of direct-beam sunlight interception at each structural scale. Total mean reductions in STAR from the leaf to the crown level were 0.39 to 0.06 in P. engelmannii and 0.46 to 0.02 in A. lasiocarpa. These reductions in STAR occurred for both species as structural scale increased due to a more upright leaf inclination, increased leaf twisting and curvature, or greater mutual shading among plant structures. A steeper leaf inclination between the leaf and shoot level accounted for 26 +/- 19% (95% C.I.) of the total leaf-to-crown STAR reduction; mutual shading among leaves on shoots caused a 14 +/- 7% reduction, whereas leaf curvature and twisting accounted for 22 +/- 3% for a total reduction of 62 +/- 8%. The STAR varied slightly from the shoot to the branch level (+/- 7%) except for a 26% reduction in shade shoots of A. lasiocarpa as a result of increased mutual shading among leaves at lateral nodes. Another substantial reduction in STAR occurred from the branch to the crown level (35 +/- 3% of total) as a result of shading of one branch layer by another within the crown. Thus, light interception efficiency decreased as structural scale increased in both species, especially from the leaf to the shoot level and from the branch to the crown level.     

Evaluating changes in switchgrass physiology,biomass, and light-use efficiency under artificial shade to estimate yields if intercropped with Pinus taeda L.

There is growing interest in using switchgrass (Panicum virgatum L.) as a biofuel intercrop in forestry systems. However, there are limited data on the longevity of intercropped bioenergy crops, particularly with respect to light availability as the overstory tree canopy matures. Therefore, we conducted a greenhouse study to determine the effects of shading on switchgrass growth. Four treatments, each with different photosynthetically active radiation (PAR) levels, were investigated inside the greenhouse: control (no shade cloth, 49 % of full sunlight), low (under 36 % shade cloth), medium (under 52 % shade cloth), and heavy shade (under 78 % shade cloth). We determined the effect of shading from March to October 2011 on individually potted, multi-tillered switchgrass transplants cut to a stubble height of 10 cm. In the greenhouse, there was a reduction in tiller number, tiller height, gas exchange rates (photosynthesis and stomatal conductance), leaf area, above- and belowground biomass and light-use efficiency with increasing shade. Total (above- and belowground) biomass in the control measured 374 ± 22 compared to 9 ± 2 g pot^?1 under heavy shade (11 % of full sunlight). Corresponding light-use efficiencies were 3.7 ± 0.2 and 1.4 ± 0.2 g MJ^?1, respectively. We also compared PAR levels and associated aboveground switchgrass biomass from inside the greenhouse to PAR levels in the inter-row regions of a range of loblolly pine (Pinus taeda L.) stands from across the southeastern United States (U.S.) to estimate when light may limit the growth of intercropped species under field conditions. Results from the light environment of loblolly pine plantations in the field suggest that switchgrass biomass will be significantly reduced at a loblolly pine leaf area index between 1.95 and 2.25, which occurs on average between ages 6 and 8 years across the U.S. Southeast in intensively managed pine plantations. These leaf area indices correspond to a 60–65 % reduction in PAR from open sky.     

Nutritive quality and morphological development under partial shade of some forage species with agroforestry potential

The shade environment produced in agroforestry practices affects the morphology, anatomy and chemical composition of intercropped forages and, therefore, may affect forage quality. During the summer-fall growing season of 1994 and 1995, 30 forage cultivars were grown in 7.6 L (two gallon) pots in full sun, 50%, and 80% shade created by placing shade cloth over a greenhouse frame. Fifteen legumes and grasses that were similar in yield under full sun and shade treatments were selected for quality analysis. With the exception of 'Kobe' lespedeza, D. paniculatum and D. canescens, acid detergent fiber (ADF) was either not affected or was slightly increased by shade. Except for Kobe lespedeza and 'Martin' tall fescue, shade either did not affect or slightly increased the neutral detergent fiber (NDF) of the shade-tolerant forages. Crude protein (CP) percentage increased in most shade-grown forages; however, shade had less effect on CP of legumes than of grasses. Other than Kobe lespedeza, total mass of CP per pot (TCP) was unaffected or increased under 50% shade for all species. Total crude protein of hog peanut, D. esmodium paniculatum and D. canescens was greater under both 50% shade and 80% shade than in full sun. The decrease in ADF of Kobe lespedeza, D. paniculatum and D. canescens under the shade treatments might be associated with maintenance of a higher leaf:stem weight ratio under shade. However, the leaf:stem ratios of the same cultivars grown at different light intensities did not strongly correlate with NDF, CP, and TCP values. In general, on selected species, internodal length and leaf area increased while specific leaf dry weight decreased for plants grown in shade compared to those grown in full sun. The data indicated that forage quality of some species could be enhanced by shade. This revised version was published online in June 2006 with corrections to the Cover Date.     

异质光环境中旱柳的光截取和利用反应

比较了不同光环境中旱柳枝，叶的生物量分配，形态特征，光能利用和水分特征。（1）阳性枝的枝重比显著大于全不遮荫枝；阴生枝的枝重比显著小于全部遮荫枝，而叶重比恰好相反。（2）阳生枝与全不遮荫枝的形态特征无显著差异。阴生枝的叶面积比率，比枝长和单位枝长叶数都显著大于全部遮荫枝，因而阴生枝的截光潜力大于全部遮荫枝。（3）阳性枝和阴生枝的植冠生产力指数分别显著大于全不遮荫枝和全部遮荫枝，因此，异质光环境中旱柳植株对光资源的利用效率大于同质光环境，（4）阳生枝的水分饱和亏缺和失水系数都显著小于全不遮荫枝，这暗示，前者的抗旱保水性大于后者。