Fruit load and canopy shading affect leaf characteristics and net gas exchange of 'Spring' navel orange trees

Five-year-old 'Spring' navel (Citrus sinensis (L.) Osbeck) orange trees were completely defruited, 50% defruited or left fully laden to study effects of fruit load on concentrations of nitrogen (N) and carbohydrate, net assimilation of CO2 (Ac) and stomatal conductance (gs) of mature leaves on clear winter days just before fruit harvest. Leaves on defruited trees were larger, had higher starch concentrations and greater leaf dry mass per area (LDMa) than leaves on fruited trees. Both Ac and gs were more than 40% lower in sunlit leaves on defruited trees than in sunlit leaves on trees with fruit. Leaves immediately adjacent to fruit were smaller, had lower leaf nitrogen and carbohydrate concentrations, lower LDMa and lower Ac than leaves on non-fruiting branches of the same trees. Removing half the crop increased individual fruit mass, but reduced fruit color development. Half the trees were shaded with 50% shade cloth for 4 months before harvest to determine the effects of lower leaf temperature (Tl) and leaf-to-air vapor pressure difference on leaf responses. On relatively warm days when sunlit Tl > 25 degrees C, shade increased Ac and gs, but had no effect on the ratio of internal to ambient CO2 (Ci/Ca) concentration in leaves, implying that high mesophyll temperatures in sunlit leaves were more important than gs in limiting Ac. Sunlit leaves were more photoinhibited than shaded leaves on cooler days when Tl < 25 degrees C. Shade decreased total soluble sugar concentrations in leaves, but had no effect on leaf starch concentrations. Shading had no effects on canopy volume, yield or fruit size, but shaded fruit developed better external color than sun-exposed fruit. Overall, the presence of a normal fruit crop resulted in lower foliar carbohydrate concentrations and higher Ac compared with defruited trees, except on warm days when Ac was reduced by high leaf temperatures.     

Growth,dry matter production,phenotypic plasticity,and nutritive value of three natural populations of Dactylis glomerata L. under various shading treatments

Dactylis glomerata L. is a widespread perennial grass species, which has been reported to be adapted to shaded conditions. Its populations thrive in a variety of environments. However, little information is available concerning the comparative response of its natural populations from contrasting environments under the reduced light intensity conditions that exist in silvopastoral systems. The objective of the present study was to estimate the comparative ability of three populations of D. glomerata from northern, central and southern Greece to grow under full sun, 60 % shade and 90 % shade in terms of their growth characteristics, phenotypic plasticity, dry matter production and nutritive value. Shade reduced tillering and dry matter production, increased tiller height and modified leaf characteristics. Under shade fewer leaves were grown simultaneously on the same tiller, but these were longer and thinner compared to full sun. Differentiation in response to shade among the populations examined of D. glomerata was observed mainly for leaf characteristics. The population from Pertouli (central Greece) responded better, particularly to moderate shade as it had a higher leaf area, longer leaf and higher dry matter production, compared to the others. Evidence for adaptive phenotypic plasticity to moderate shade was suggested only for this population. Additionally, Pertouli had higher nutritive value compared to Taxiarchis (northern Greece) and Crete (southern Greece) under shade. The divergent responses of natural populations of D. glomerata could justify breeding germplasm with enhanced shade tolerance.     

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.     

不同单株负载量对赤霞珠葡萄生长及果实品质的影响

为了探索不同的单株负载量对酿酒葡萄生长及果实品质的影响,以赤霞珠酿酒葡萄为试材,设置了宁夏贺兰山东麓地区砾质沙土和灌淤土上赤霞珠不同单株负载量试验,测定了不同负载量水平下葡萄果实的单果质量、可溶性总糖含量、可滴定酸含量以及叶片的营养元素和百叶鲜干质量。结果表明:砾质沙土和灌淤土上赤霞珠叶片的百叶鲜干比均是先减小后增大;2种土壤上赤霞珠葡萄叶片和叶柄中氮、磷、钾含量均是随着单株负载量的增加而减少,砾质沙土叶柄钾含量的变异幅度较大,单株负载5串的处理比单株负载25串的处理叶柄钾含量高出87.31%;砾质沙土上6年生赤霞珠单株负载15串时可溶性总糖含量达到了23.33%,单果质量最小(1.36 g),单穗质量最大(228.3 g);灌淤土上6年生赤霞珠则是单株负载10串时单果质量最小(1.35 g),可溶性总糖含量适中(17.94%),糖酸比24.24。合理的负载水平对酿酒葡萄果实品质影响较为显著,砾质沙土6年生赤霞珠合理的负载量为单株15串,灌淤土6年生赤霞珠合理的负载量为单株10串。     

Effect of hydropriming and acclimation treatments on <Emphasis Type="Italic">Quercus rugosa</Emphasis> acorns and seedlings

To facilitate restoration of the lava field forests surrounding Mexico City, we developed methods to improve the germination and field seedling performance of Quercus rugosa using hydropriming (regulated hydration of seeds in water), and we used special watering regimes to improve seedling acclimation. The size, dry mass, fresh mass and water content of seeds were measured, and curves were generated to evaluate acorn hydration and dehydration. The effects of stratification (5°C), heat shock (50°C) and scarification on germination were tested. All treated seeds and controls were germinated in control chambers at 21°C. One hydropriming cycle (PC) consisted of two hydration days followed by two dehydration days; treatments of 1, 2 and 3 PCs were tested. Seedlings from 1PC to 2PC were acclimated in a shade house under high and low watering regimes (400 and 200 mL week^?1, respectively). In the shade house and field, the effects of hydropriming and watering treatments were evaluated by measuring length, basal diameter, crown cover, number of leaves and branches and leaf area of seedlings. Dry and fresh mass were used to calculate acorn water content. Dehydration and hydration curves displayed hysteresis. Acorns exhibited physiological dormancy, which could be overcome by stratification or by 1 month of storage. 1PC led to increased germination rates and final germination. In both the shade house and field, 1PC showed a positive effect on all seedling growth parameters except branch number. Field survival was not affected. Generally, 1PC favoured efficient seed germination, seedling vigour and homogenous plant production.     

Interactive effects of shade and irrigation on the performance of seedlings of three Mediterranean Quercus species

Shade and irrigation are frequently used to increase the success of Mediterranean Quercus spp. plantations. However, there is controversy about the combined effects of these treatments on plant performance. We assessed the effects of two irradiances (full sunlight and moderate shade) and two summer watering regimes (high (daily) and low (alternate days)) on leaf and whole-plant traits of 1-year-old seedlings of Quercus coccifera, Q. ilex subsp. ballota and Q. faginea grown outdoors for 8.5 months. Leaf traits included measures of morphology, nitrogen concentration, gas exchange and photochemical efficiency, and measures of whole-plant traits included biomass allocation patterns, growth phenology, across-summer leaf area change and relative growth rate (RGR). Moderate shade reduced leaf mass per area, increased photochemical efficiency, maximum carbon assimilation rate (Amax) and allocation to leaves, and prolonged the growing period in one or more of the species. Daily watering in summer increased Amax of Q. ilex and prolonged the growing period of Q. ilex and Q. faginea. Both treatments tended to increase RGR. The effect of shade was greater in the low-watering regime than in the high-watering regime for two of the 15 studied traits, with treatment effects being independent for the remaining 13 traits. Leaf nitrogen and the ability to maintain leaf area after the arid period, rather than biomass allocation traits, explained the variation in seedling RGR. Trait responsiveness to the treatments was low and similar among species and between study scales, being unexpectedly low in Q. faginea leaves. This may be because selective pressures on leaf plasticity act differently in deciduous and evergreen species. We conclude that moderate shade and daily summer watering enhance the performance of Mediterranean Quercus seedlings through species-specific mechanisms.     

Ecophysiological and morphological responses to shade and drought in two contrasting ecotypes of Prunus serotina

Photosynthesis (A), water relations and stomatal reactivity during drought, and leaf morphology were evaluated on 2-year-old, sun- and shade-grown Prunus serotina Ehrh. seedlings of a mesic Pennsylvania seed source and a more xeric Wisconsin source. Wisconsin plants maintained higher A and leaf conductance (g(wv)) than Pennsylvania plants during the entire drought under sun conditions, and during the mid stages of drought under shade conditions. Compared to shade plants, sun plants of both sources exhibited a more rapid decrease in A or % A(max) with decreasing leaf water potential (Psi). Tissue water relations parameters were generally not significantly different between seed sources. However, osmotic potentials were lower in sun than shade plants under well-watered conditions. Following drought, shade plants, but not sun plants, exhibited significant osmotic adjustment. Sun leaves had greater thickness, specific mass, area and stomatal density and lower guard cell length than shade leaves in one or both sources. Wisconsin sun leaves were seemingly more xerophytic with greater thickness, specific mass, and guard cell length than Pennsylvania sun leaves. No source differences in leaf structure were exhibited in shade plants. Stomatal reactivity to sun-shade cycles was similar between ecotypes. However, well-watered and droughted plants differed in stomatal reactivity within and between multiple sun-shade cycles. The observed ecotypic and phenotypic variations in ecophysiology and morphology are consistent with the ability of Prunus serotina to survive in greatly contrasting environments.     

Fruit load and branch ring-barking affect carbon allocation and photosynthesis of leaf and fruit of Coffea arabica in the field

Increasing fruit load (from no berries present to 25, 50 and 100% of the initial fruit load) significantly decreased branch growth on 5-year-old coffee (Coffea arabica L.) trees of the dwarf cultivar 'Costa Rica 95', during their third production cycle. Ring-barking the branches further reduced their growth. Berry dry mass at harvest was significantly reduced by increasing fruit load. Dry matter allocation to berries was four times that allocated to branch growth during the cycle. Branch dieback and berry drop were significantly higher at greater fruit loads. This illustrates the importance of berry sink strength and indicates that there is competition for carbohydrates between berries and shoots and also among berries. Leaf net photosynthesis (P(n)) increased with increasing fruit load. Furthermore, leaves of non-isolated branches bearing full fruit load achieved three times higher P(n) than leaves of isolated (ring-barked) branches without berries, indicating strong relief of leaf P(n) inhibition by carbohydrate demand from berries and other parts of the coffee tree when excess photoassimilates could be exported. Leaf P(n) was significantly higher in the morning than later during the day. This reduction in leaf P(n) is generally attributed to stomatal closure in response to high irradiance, temperature and vapor pressure deficit in the middle of the day; however, it could also be a feedback effect of reserves accumulating during the morning when climatic conditions for leaf P(n) were optimal, because increased leaf mass ratio was observed in leaves of ring-barked branches with low or no fruit loads. Rates of CO(2) emission by berries decreased and calculated photosynthetic rates of berries increased with increasing photosynthetic photon flux (PPF) especially at low PPFs (0 to 100 micromol m(-2) s(-1)). The photosynthetic contribution of berries at the bean-filling stage was estimated to be about 30% of their daily respiration costs and 12% of their total carbon requirements at PPF values commonly experienced in the field (200 to 500 micromol m(-2) s(-1)).     

Leaf optical properties in Venezuelan cloud forest trees

Leaf optical properties and related leaf characteristics were compared for thirteen cloud forest tree species differing in successional status. Sun leaves were sampled for the eight pioneer species and sun and shade leaves were sampled for the five climax species. Sun leaves had a slightly higher absorptance than shade leaves, although differences were small. Sun leaves had a higher leaf mass per unit area (LMA) and a lower chlorophyll concentration per unit leaf mass, resulting in similar chlorophyll concentrations per unit leaf area and hence similar light harvesting capacities as shade leaves. However, shade leaves realized a higher efficiency of absorptance per unit leaf biomass than sun leaves. There were few differences in leaf characteristics of sun leaves between the climax and pioneer species. Absorptance values of cloud forest species were comparable with values reported for rain forest and more seasonal forest species. Intraspecific variation in leaf absorptance was largely the result of variation in LMA, whereas interspecific variation in leaf absorptance was largely a result of variation in chlorophyll concentration per unit leaf area.     

Effects of CO2 enrichment on the photosynthetic light response of sun and shade leaves of canopy sweetgum (Liquidambar styraciflua) in a forest ecosystem

To investigate whether sun and shade leaves respond differently to CO2 enrichment, we examined photosynthetic light response of sun and shade leaves in canopy sweetgum (Liquidambar styraciflua L.) trees growing at ambient and elevated (ambient + 200 microliters per liter) atmospheric CO2 in the Brookhaven National Laboratory/Duke University Free Air CO2 Enrichment (FACE) experiment. The sweetgum trees were naturally established in a 15-year-old forest dominated by loblolly pine (Pinus taeda L.). Measurements were made in early June and late August 1997 during the first full year of CO2 fumigation in the Duke Forest FACE experiment. Sun leaves had a 68% greater leaf mass per unit area, 63% more leaf N per unit leaf area, 27% more chlorophyll per unit leaf area and 77% greater light-saturated photosynthetic rates than shade leaves. Elevated CO2 strongly stimulated light-saturated photosynthetic rates of sun and shade leaves in June and August; however, the relative photosynthetic enhancement by elevated CO2 for sun leaves was more than double the relative enhancement of shade leaves. Elevated CO2 stimulated apparent quantum yield by 30%, but there was no interaction between CO2 and leaf position. Daytime leaf-level carbon gain extrapolated from photosynthetic light response curves indicated that sun leaves were enhanced 98% by elevated CO2, whereas shade leaves were enhanced 41%. Elevated CO2 did not significantly affect leaf N per unit area in sun or shade leaves during either measurement period. Thus, the greater CO2 enhancement of light-saturated photosynthesis in sun leaves than in shade leaves was probably a result of a greater amount of nitrogen per unit leaf area in sun leaves. A full understanding of the effects of increasing atmospheric CO2 concentrations on forest ecosystems must take account of the complex nature of the light environment through the canopy and how light interacts with CO2 to affect photosynthesis.     

Effects of shade on growth, biomass allocation and leaf morphology in European yew (Taxus baccata L.)

The impact of shade on the growth of European yew (Taxus baccata L.) saplings was investigated over a three-year period using artificial shading to simulate four different light regimes (3, 7, 27 and 100 % relative photosynthetic photon flux density, RPPFD). There was no mortality attributable to shading even under the 3 % RPPFD treatment. Increasing shade was positively associated with specific leaf area, leaf length, leaf width and total chlorophyll content, but negatively associated with plant height, stem diameter, total dry weight and root to leaf and shoot ratio. Discoloration of the foliage occurred in plants grown in 100 % RPPFD conditions (resulting in reduced growth rates) and those transferred to 100 % RPPFD conditions after being shade-acclimated for 2 years. Evidence suggests that T. baccata has the ability to regenerate beneath a lighter canopy but beneath denser canopies gap dynamics will play an important role in facilitating successful regeneration and this needs to be reflected in management of natural populations of this declining species.     

文冠果表型性状的相关性及主成分分析

文冠果种子产量低,这是制约其大力开发与利用的瓶颈。为给文冠果优良品种的选育和文冠果资源评价和利用提供科学依据,对新疆维吾尔自治区奇台县的160株文冠果单株的种子质量、结果个数、果实质量等21个性状指标进行了调查,并采用变异分析、相关性分析和主成分分析等分析方法,对影响其产量的主要因素进行了探讨。变异分析结果表明:被调查的160株文冠果单株各性状指标间均存在明显差异;其中,单株结果个数的变异系数最大,为82.98%;叶密度的变异系数最小,为8.77%。相关性分析结果表明:单株种子质量与树高、地径、冠幅、果实纵径、种子长、单果种子质量、单株结果个数、单株果实质量、百果质量、百粒种子质量、叶长、叶宽、叶面积、果皮厚度、单果种子粒数及种子宽之间均存在显著正相关。主成分分析结果表明:从21个性状指标中提取了7个主成分,即单果产量、单株产量、果实产量、叶形、种形、果形、树形这7个因子,其累积贡献率达74.693%;依据各主成分贡献率的大小,在育种中可选择单果种子质量大、单株结果个数多、单株果实质量大、叶长、种子长、果纵径长、冠幅大的单株作为优良高产植株。     

Differences in leaf functional traits between red and green leaves of two evergreen shrubs <Emphasis Type="Italic">Photinia</Emphasis> × <Emphasis Type="Italic">fraseri</Emphasis> and <Emphasis Type="Italic">Osmanthus fragrans</Emphasis>

Leaf functional traits are adaptations that enable plants to live under different environmental conditions. This study aims to evaluate the differences in leaf functional traits between red and green leaves of two evergreen shrubs Photinia × fraseri and Osmanthus fragrans. Specific areas of red leaves are higher than that of green leaves in both species. Thus, the material investment per unit area and per lamina of red leaves is significantly lower than that of green leaves, implying an utmost effort of red leaves to increase light capture and use efficiency because of their low leaf-chlorophyll concentration. The higher petiole length of green leaves compared with that of red leaves indicates that adult green leaves may have large fractional biomass allocation to support the lamina structures in capturing light with maximum efficiency and obtaining a high growth rate. The high range of the phenotypic plasticity of leaf size, leaf thickness, single-leaf wet and dry weights, and leaf moisture of green leaves may be beneficial in achieving efficient control of water loss and nutrient deprivation. The high range of phenotypic plasticity of leaf chlorophyll concentration of red leaves may be advantageous in increasing resource (especially light) capture and use efficiency because this leaf type is juvenile in the growth stage and has low leaf-chlorophyll concentration.     

Photosynthetic acclimation of overstory Populus tremuloides and understory Acer saccharum to elevated atmospheric CO2 concentration: interactions with shade and soil nitrogen

We exposed Populus tremuloides Michx. and Acer saccharum Marsh. to a factorial combination of ambient and elevated atmospheric CO2 concentrations ([CO2]) and high-nitrogen (N) and low-N soil treatments in open-top chambers for 3 years. Our objective was to compare photosynthetic acclimation to elevated [CO2] between species of contrasting shade tolerance, and to determine if soil N or shading modify the acclimation response. Sun and shade leaf responses to elevated [CO2] and soil N were compared between upper and lower canopy leaves of P. tremuloides and between A. saccharum seedlings grown with and without shading by P. tremuloides. Both species had higher leaf N concentrations and photosynthetic rates in high-N soil than in low-N soil, and these characteristics were higher for P. tremuloides than for A. saccharum. Electron transport capacity (Jmax) and carboxylation capacity (Vcmax) generally decreased with atmospheric CO2 enrichment in all 3 years of the experiment, but there was no evidence that elevated [CO2] altered the relationship between them. On a leaf area basis, both Jmax and Vcmax acclimated to elevated [CO2] more strongly in shade leaves than in sun leaves of P. tremuloides. However, the apparent [CO2] x shade interaction was largely driven by differences in specific leaf area (m2 g-1) between sun and shade leaves. In A. saccharum, photosynthesis acclimated more strongly to elevated [CO2] in sun leaves than in shade leaves on both leaf area and mass bases. We conclude that trees rooted freely in the ground can exhibit photosynthetic acclimation to elevated [CO2], and the response may be modified by light environment. The hypothesis that photosynthesis acclimates more completely to elevated [CO2] in shade-tolerant species than in shade-intolerant species was not supported.     

Tradeoff between shade adaptation and mitigation of photoinhibition in leaves of Quercus mongolica and Acer mono acclimated to deep shade

We investigated susceptibility to photoinhibition in leaves acclimated to different light regimes in intermediately shade-tolerant Japanese oak (Quercus mongolica Fisch. ex Turcz. var. crispula (Blume) Ohashi) and shade-tolerant Japanese maple (Acer mono Maxim. var. glabrum (Lév. et Van't.) Hara), to elucidate adaptability to gap formation in leaves differing in shade acclimation. We hypothesized that there is a tradeoff between shade adaptation and capacity to mitigate photoinhibition associated with leaf morphology. We simultaneously measured chlorophyll fluorescence and gas exchange in seedlings that had been grown in full sunlight (open), 10% of full sun (moderate shade) and 5% of full sun (deep shade). Shade-tolerant A. mono adapted to deep shade through changes in leaf morphology, lowering its leaf mass per area (LMA), but Q. mongolica showed little change in LMA between moderate and deep shade. Photochemical quenching (qP) did not differ between species in full sunlight and moderate shade; however, in deep shade, qP of Q. mongolica was higher than that of A. mono, suggesting that Q. mongolica grown in deep shade is less susceptible to photoinhibition at gap formation. This is consistent with the finding that chronic photoinhibition 3 days after the transfer to full sunlight, indicated by the decrease in maximum photochemical efficiency, Fv/Fm, at predawn, was less in deep-shade-grown Q. mongolica than in deep shade-grown A. mono. In deep shade, the electron transport rate (ETR) of Q. mongolica was higher than that of A. mono, whereas thermal energy dissipation through photosystem II antennae, indicated by non-photochemical quenching, was lower in Q. mongolica than in A. mono. In deep shade, the greater ETR capacity in Q. mongolica in association with higher LMA and higher leaf N content could contribute to maintaining high qP and mitigating photoinhibition. These results indicate that, by maintaining a high electron transport capacity even in deep shade, the gap-dependent and intermediate-shade-tolerant Q. mongolica trades improved shade adaptation for higher growth potential when a gap event occurs.     

Influence of irradiance on photosynthesis, morphology and growth of mangosteen (Garcinia mangostana L.) seedlings

The influence of shading intensity on growth, morphology and leaf gas exchange of mangosteen (Garcinia mangostana L.) seedlings was investigated over a 2-year period. Diurnal gas exchange studies revealed significantly higher carbon gain for leaves grown in 20 or 50% shade compared to leaves grown in 80% shade. Seedlings grown in 20 or 50% shade accumulated significantly more dry weight than seedlings grown in 80% shade during the 2-year study period. Seedlings grown in decreased shade showed decreased leaf size, increased leaf thickness, lower specific leaf area (SLA) and higher stomatal frequency. Less shaded seedlings also allocated relatively more dry matter to roots than shaded seedlings and exhibited a significant reduction in leaf area relative to total plant dry weight (leaf area ratio). Increased leaf number, enhanced branching and shorter internodes resulted in a more compact appearance of less shaded seedlings. Irrespective of light conditions, mangosteen seedlings exhibited inherently slow growth because of low photosynthetic rates per unit leaf area, low SLA, low leaf area ratios and inefficient root systems.     

Evaluation of leaf display of evergreen broadleaved tree species and deciduous tree species in warm temperate conifer plantations

We investigated the sapling leaf display in the shade among trees of various leaf lifespans co-occurring under the canopy of a warm-temperate conifer plantation. We measured leaf-area ratio (aLAR) and morphological traits of saplings of evergreen broadleaved tree species and a deciduous tree species. Although we found large interspecific and intraspecific differences in aLAR even among saplings of similar size in the homogeneous light environment, we did not find a consistent trend in aLAR with leaf lifespan among the species. While deciduous trees annually produced a large leaf area, some evergreen broadleaved trees retained their leaves across years and had aLAR values as high as those of deciduous trees. Among leaf-level, shoot-level, and individual-level morphological traits, aLAR was positively correlated with current-year shoots mass per aboveground biomass in deciduous trees, and with the area of old leaves per aboveground mass in evergreen broadleaved trees. Thus, tree-to-tree variation in the degrees of annual shoot production and the accumulation of old leaves were responsible for the interspecific and intraspecific variations in aLAR.     

Growth and Development Responses of Three Acacia Species to Long-Term Light Regimes

Abstract

Three species of Acacia which are used in various forestry systems in Micronesia are the Acacia confusa Merr., A. mangium Willd., and A. auriculiformis Cunn. ex Benth. The responses of these species to limited light have not been established. The objective of this study was to determine the growth responses of these three species to light ranging from 19% to 100% sunlight transmission in order to determine the relative tolerance of these species to shade. Several common adaptations to low light occurred with all three species, such as increased leaf surface area in relation to mass of supportive tissue and reduced leaf mass per unit leaf area. However, other adaptations to low light were not evident. Numerous indicators of growth were decreased at low levels of sunlight transmission. The relative reduction in growth at 19% transmission compared with 100% transmission was similar for each species. The results indicate that these Acacia species exhibit some adaptations to development in limited light, but they are not tolerant of shade. All three species are thus classified as shade avoiders, and have limited application under shaded conditions in any Pacific island forestry system.     

Leaf litter and its nutrient contribution in the cacao agroforestry system

Cacao agroforestry systems (cacao-AFS) produce abundant litter. After decomposing, litter releases nutrients into the soil. The aim of this research was to estimate litter production and its nutrient content in 35- and 55-year-old cacao-AFS. The research was conducted in three cacao-AFS of each age, in Cardenas, Tabasco, México. Four traps per cacao-AFS were used to collect litter. Litter was collected every 15 days for one year. It was then fractioned into cacao leaves, shade tree leaves, petioles, branches and stems, and cacao flowers and fruits. To determine nutrient content of litter, samples were composited by age of cacao-AFS and by season of the year. Then chemical analysis was done in triplicate. Data were subjected to analysis of variance, orthogonal contrasts, and Student t and Duncan tests. Cacao-AFS produce litter all year. Thirty-five-year-old cacao-AFS produced more litter than 55-year-old cacao-AFS (2042 vs 1570 kg DM ha^?1 year^?1). Except for the shade tree leaf fraction (559.5 vs 642 kg DM ha^?1), 35-year-old cacao-AFS were superior to 55-year-old cacao-AFS in all the other litter fractions. Cacao leaf fraction was the main source of litter in cacao-AFS of both ages. Neither age of cacao-AFS nor the season of the year affected N, K, Zn or S content in litter. Orthogonal contrasts indicated statistical differences between ages of cacao-AFS for P, Ca, and Fe content in litter. Both N–P–K–Ca–Mg contents in litter of 35-year-old cacao-AFS (1.2–0.4–1.2–1.7–0.4%) and in litter of 55-year-old cacao-AFS (1.1–0.6–1.2–1.4–0.4%) are enough to recover the nutrients extracted by the cacao crop.     

Shade acclimation in the forage grass Festuca Pallescens: biomass allocation and foliage orientation

Plants can acclimate to shade through different processes. In particular, they can modify their biomass allocation and the architecture in order to increase light interception. The objective of this study was to evaluate the shade acclimation capacity of Festuca pallescens (St. Ives) Parodi, as part of research concerning the use of this species in silvopastoral systems in Patagonia, Argentina. Biomass allocation was estimated from the leaf and root dry weights of plants growing in an open pasture and forested plots. Crown architecture of plants growing in the open and in two shade treatments was studied dividing each plant in three concentric cylinders, within which leaf angles and leaf area were measured. Light interception of plants in each treatment was estimated from the projected leaf areas and the relative amount of radiation reaching each location. Biomass allocation changed significantly in plants growing under shade conditions, increasing the proportion of leaves relative to the roots (Leaf Mass Fraction = 0.29 (SD: 0.12) and 0.40 (SD: 0.09) in plants in the open and under shade, respectively). Also, mean leaf inclination angles changed in plants growing under shade conditions, allowing an increase in light interception of approximately 35% compared to plants with the crown architecture typical of the open treatment. Previous studies have shown that F. pallescens does not change its photosynthetic response to light under shade conditions. Therefore, we conclude that the reported changes in biomass allocation and crown architecture, in addition to the increment in specific leaf area explain the relatively high shade tolerance of this species. This revised version was published online in June 2006 with corrections to the Cover Date.