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.     

Shade tolerance, photoinhibition sensitivity and phenotypic plasticity of Ilex aquifolium in continental Mediterranean sites

Shade tolerance, plastic phenotypic response to light and sensitivity to photoinhibition were studied in holly (Ilex aquifolium L.) seedlings transported from the field to a greenhouse and in adult trees in the field. All plants were growing in, or originated from, continental Mediterranean sites in central Spain. Seedlings tolerated moderate but not deep shade. Mortality was high and growth reduced in 1% sunlight. Survival was maximal in 12% sunlight and minimal in full sunlight, although the relative growth rate of the seedlings surviving in high light was similar to that of plants in moderate shade. Maximum photochemical efficiency at predawn was significantly lower in sun plants than in shade plants in the field, revealing chronic photoinhibition that was most pronounced in winter. Plasticity in response to available light varied according to the variable studied, being low for photosynthetic capacity and stomatal conductance, and high for specific leaf area, root:shoot ratio and leaf area ratio, particularly in seedlings. Differences in water relations and hydraulic features between sun and shade plants in the field were marginal. High water potential at the turgor loss point of field-grown plants suggested that holly is sensitive to drought during both the seedling and the adult stage. Low relative growth rates in both high and low light with low physiological plasticity in response to light indicate the existence of a stress-tolerance mechanism. We conclude that holly is a facultative understory plant in areas of oceanic and relatively mild climate, but an obligate understory plant in dry continental areas such as the study site. The impact of abandonment of traditional management practices and climate change on these Mediterranean populations is discussed.     

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.     

Induction of photosynthesis and importance of limitations during the induction phase in sun and shade leaves of five ecologically contrasting tree species from the temperate zone

We examined the principal differences in photosynthetic characteristics between sun and shade foliage and determined the relative importance of biochemical and stomatal limitations during photosynthetic induction. Temperate-zone broadleaf and conifer tree species, ranging widely in shade tolerance, were investigated from one locality in the Czech Republic. The study species included strongly shade-tolerant Abies alba Mill. and Tilia cordata Mill., less shade-tolerant Fagus sylvatica L. and Acer pseudoplatanus L. and sun-demanding Picea abies (L.) Karst. In the fully activated photosynthetic state, sun foliage of all species had significantly higher maximum CO(2) assimilation rates, maximum stomatal conductance and maximum rates of carboxylation than shade foliage. Compared with shade leaves, sun leaves had significantly higher nocturnal stomatal conductances. In all species, shade foliage tended to have higher induction states 60 s after leaf illumination than sun foliage. Sun and shade foliage did not differ in the rate of disappearance of the transient biochemical limitation during the induction phase. Longer time periods were required to reach 90% photosynthetic induction and 90% stomatal induction in sun foliage than in shade foliage of the less shade-tolerant F. sylvatica and A. pseudoplatanus and in sun-demanding P. abies; however, in sun foliage of the strongly shade-tolerant species T. cordata and A. alba, the time needed for photosynthetic induction was similar to, or less than, that for shade foliage. Shade but not sun needles of P. abies and A. alba had significantly slower induction kinetics than the broadleaf tree species. Among species, the sun-demanding P. abies exhibited the shortest stomatal induction times in both sun and shade leaves. Independently of shade tolerance ranking, the transient stomatal and total limitations that characterize photosynthetic induction were relieved significantly earlier in shade foliage than in sun foliage. Sun foliage generally exhibited a hyperbolic photosynthetic induction response, whereas a sigmoidal induction response was more frequent in shade foliage. The different relative proportions of transient biochemical and stomatal limitations during photosynthetic induction in sun and shade foliage indicate an essential role of stomata in photosynthetic limitation during induction, mainly in shade foliage, with a consequent influence on the shape of the photosynthetic induction curve.     

Growth,photosynthetic performance and shade adaptation of rubber (Hevea brasiliensis) grown in natural shade

We compared growth, photosynthetic performance and shade adaptation of rubber (Hevea brasiliensis Muell. Arg.) plants growing in natural shade (33, 55 and 77% reduction in incoming radiation) to control plants growing in full sunlight. Stem diameter and plant height, measured over a 15-month period, were greatest in plants grown in full sunlight, and both parameters decreased with increasing shade. At 7 and 14 months after planting (MAP), total plant dry mass was highest in control plants and lowest in plants in 77% shade. Expansion of the fourth leaf whorl, monitored at 5-6 MAP, was slowest in plants in 77% shade and fastest in unshaded plants, which had more leaves and higher leaf areas and inter-whorl shoot lengths. In response to increasing shade, specific leaf area increased, whereas leaf weight ratio and relative growth rate decreased. Chlorophyll a/b ratio decreased with increasing shade, indicating shade-induced partitioning of chlorophyll into light-harvesting complexes. Compared to the response in unshaded plants, CO2 assimilation saturated at lower photosynthetic photon flux densities in plants in 77% shade, with a lower upper-asymptote to the light response curve. Chlorophyll fluorescence revealed no evidence of sustained photoinhibitory damage in unshaded plants. Dynamic photoinhibition decreased with increasing shade, with the greatest depression in the ratio of variable to maximal fluorescence around midday. We conclude that shade adaptation and shade-induced reductions in dynamic photoinhibition account for the enhanced early growth of rubber in light shade.     

遮荫下2种热带树苗叶片光合特性和抗氧化酶系统对自然降温的响应

选取 2种遮荫条件下 (8%、2 5 %全光照 )西双版纳地区热带雨林上层树种滇南红厚壳和中层树种玉蕊的幼苗 ,测定了雨季末自然降温过程中 2种植物叶片光合特性、抗氧化酶 (SOD、CAT、APX)活性、丙二醛 (MDA)含量的变化。研究结果表明 :低温使 2种植物单位面积叶绿素含量和净光合速率明显下降 ,生长在中等光强下的幼苗比低光强下的幼苗下降的比率大 ,滇南红厚壳比玉蕊下降的比率大。在降温初期 ,即从 10月到 12月初 ,2种光强下 2种植物SOD和APX活性上升 ,CAT活性出现波动 ,黎明光化学效率和MDA含量基本没有变化 ,光合机构没有受到氧化破坏。 12月末 ,中等光强下滇南红厚壳黎明光化学效率降低 ,SOD和APX活性有所下降 ,MDA含量显著上升 ;而低光强下的滇南红厚壳和 2种光强下玉蕊SOD和APX活性仍在上升 ,MDA含量变化较小。这表明 ,2种光强下 2种热带树苗是以不同的方式响应自然降温 ,低温对中等光强下的植物比低光强下的影响大 ,上层树种滇南红厚壳受到的影响比中层树种玉蕊大。同时 ,根据降温过程中抗氧化酶活性和MDA含量的变化 ,大致可将西双版纳热带雨林幼苗对低温的响应分为 2个阶段 ,即第 1阶段为低温适应阶段 ,抗氧化酶活性上升 ,光合机构没有受到明显的氧化破坏 ;第 2阶段 ,中等光强下滇南红厚壳抗氧化酶     

Leaf senescence and late-season net photosynthesis of sun and shade leaves of overstory sweetgum (Liquidambar styraciflua) grown in elevated and ambient carbon dioxide concentrations

We examined the effects of elevated CO2 concentration ([CO2]) on leaf demography, late-season photosynthesis and leaf N resorption of overstory sweetgum (Liquidambar styraciflua L.) trees in the Duke Forest Free Air CO2 Enrichment (FACE) experiment. Sun and shade leaves were subdivided into early leaves (formed in the overwintering bud) and late leaves (formed during the growing season). Overall, we found that leaf-level net photosynthetic rates were enhanced by atmospheric CO2 enrichment throughout the season until early November; however, sun leaves showed a greater response to atmospheric CO2 enrichment than shade leaves. Elevated [CO2] did not affect leaf longevity, emergence date or abscission date of sun leaves or shade leaves. Leaf number and leaf area per shoot were unaffected by CO2 treatment. A simple shoot photosynthesis model indicated that elevated [CO2] stimulated photosynthesis by 60% in sun shoots, but by only 3% in shade shoots. Whole-shoot photosynthetic rate was more than 12 times greater in sun shoots than in shade shoots. In senescent leaves, elevated [CO2] did not affect residual leaf nitrogen, and nitrogen resorption was largely unaffected by atmospheric CO2 enrichment, except for a small decrease in shade leaves. Overall, elevated [CO2] had little effect on the number of leaves per shoot at any time during the season and, therefore, did not change seasonal carbon gain by extending or shortening the growing season. Stimulation of carbon gain by atmospheric CO2 enrichment in sweetgum trees growing in the Duke Forest FACE experiment was the result of a strong stimulation of photosynthesis throughout the growing season.     

Ionic homeostasis and reactive oxygen species control in leaves and xylem sap of two poplars subjected to NaCl stress

We investigated the effects of increasing soil NaCl concentration on intracellular compartmentalization of salt and on the activities of antioxidant enzymes (superoxide dismutase (SOD), ascorbic peroxidase (APX), catalase (CAT) and glutathione reductase (GR)) and their role in the regulation of reactive oxygen species (ROS; O(2)(-*) and H(2)O(2)) in leaves and xylem sap of salt-tolerant Populus euphratica Oliv. and salt-sensitive P. popularis cv. 35-44. Mesophyll cells of P. euphratica exhibited a high capacity for NaCl exclusion and compartmentalization of salt in vacuoles compared with P. popularis. In P. popularis, the salt treatment resulted in large accumulations of Na(+) and Cl(-) in leaves that induced significant increases in O(2)(-*) and H(2)O(2) production despite marked increases in the activities of antioxidant enzymes in leaves and xylem sap. Separation of the isoforms of leaf SOD, APX and CAT by polyacrylamide gel electrophoresis followed by in-gel activity staining revealed that the salt-induced activities of APX and CAT were the result of increases in activities of all the isoenzymes. Leaf injury and shedding of aged leaves occurred following the oxidative burst in P. popularis, indicating that the increased activities of antioxidant enzymes in P. popularis were insufficient to counter the harmful effects of ROS at high soil NaCl concentrations. Unlike P. popularis plants, P. euphratica plants did not exhibit an oxidative burst in response to the NaCl treatments, because of (1) a high salt exclusion capacity and effective compartmentalization of salt in vacuoles, and (2) up-regulation of antioxidant enzymatic activities after the onset of salt stress. We conclude that P. euphratica plants subjected to saline conditions control ROS homeostasis through two pathways: (1) by maintaining cellular ionic homeostasis and thereby limiting the NaCl-induced enhancement of ROS production under long-term saline conditions; and (2) by rapidly up-regulating antioxidant defenses to prevent oxidative damage.     

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.     

Physiological responses of saplings of rainforest timber species differing in successional group to light regimes and nitrogen

Saplings of 19 valuable rain forest timber species representative of three successional status groups (early secondary, late secondary and climax) were grown in a polyhouse to examine their responses to three light intensity/quality treatments and nitrogen supply. Solar radiation was modified using painted polyethylene sheet to mimic natural light environments across a rain forest vertical column as follows: 1. Transparent plastic, 80% of full sunlight, R:FR = 0.95, 2. Blue shade, 14% of full sunlight, R:FR = 0.69; 3. Green shade, 7% of full sunlight, R:FR = 0.50. Transparent plastic conditions promoted an increase in stem height and diameter (i.e., growth), leaf thickness and gas exchange per unit leaf area. Additional nitrogen availability enhanced growth and specific leaf area (i.e., leaves were thinner), particularly in the full sun environment and on early secondary and late secondary successional species, but did not influence photosynthetic rate. Successional status of the species did not affect photosynthetic rate although early secondary successional species grew faster and had fewer branches than species of the other successional groups. We recommend that for a successful mixed stand the high-light requiring species should be planted first, with increased nitrogen supply, and the shade tolerant species should be introduced later with no extra nitrogen supply required.     

干旱和遮荫对中国沙棘水碳平衡和黄酮化合物含量的影响

目的 研究干旱和遮荫对中国沙棘生长、水碳代谢和黄酮类化合物含量的影响有助于全面了解沙棘的生态适应性,同时有利于了解干旱和遮荫对沙棘黄酮类化合物含量的影响。 方法 本研究以中国沙棘为研究对象,比较了干旱（40%土壤水分饱和含水量）、遮荫（遮光50%）及二者交互处理下的水势、枝条导水损失率（PLC）、光合作用、生物量、非结构性碳水化合物（NSC）含量和黄酮类化合物含量的变化。 结果 干旱处理显著降低了中国沙棘的枝条水势、净光合速率、叶生物量、根和茎的NSC含量,显著增加了枝条PLC、根生物量;遮荫显著降低了净光合速率、叶生物量、根茎叶的NSC含量和黄酮含量,干旱和遮荫的共同作用显著增加了枝条PLC,降低了叶净光合速率、根茎叶生物量和NSC含量;且叶黄酮类化合物含量与枝条水势呈显著负相关。 结论 干旱会影响中国沙棘的水分状况、碳摄取,降低茎和根的NSC储藏,增加黄酮类化合物含量,遮荫会影响中国沙棘的光合速率、根茎叶的NSC储藏,干旱和遮荫的共同作用进一步降低了中国沙棘的水分输导能力、光合作用、生物量积累和NSC储藏,遮荫降低了中国沙棘对干旱胁迫的适应能力,适度干旱有利于沙棘黄酮类化合物的合成。     

Effects of elevated CO(2) and light availability on the photosynthetic light response of trees of contrasting shade tolerance

Photosynthetic light response curves (A/PPFD), leaf N concentration and content, and relative leaf absorbance (alpha(r)) were measured in 1-year-old seedlings of shade-intolerant Betula papyrifera Marsh., moderately shade-tolerant Quercus rubra L. and shade-tolerant Acer rubrum L. Seedlings were grown in full sun or 26% of full sun (shade) and in ambient (350 ppm) or elevated (714 ppm) CO(2) for 80 days. In the shade treatments, 80% of the daily PPFD on cloud-free days was provided by two 30-min sun patches at midday. In Q. rubra and A. rubrum, leaf N concentration and alpha(r) were significantly higher in seedlings in the shade treatments than in the sun treatments, and leaf N concentration was lower in seedlings in the ambient CO(2) treatments than in the elevated CO(2) treatments. Changes in alpha(r) and leaf N content suggest that reapportionment of leaf N into light harvesting machinery in response to shade and elevated CO(2) tended to increase with increasing shade tolerance of the plant. Shifts induced by elevated CO(2) in the A/PPFD relationship in sun plants were largest in B. papyrifera and least in A. rubrum: the reverse was true for shade plants. Elevated CO(2) resulted in increased light-saturated A in every species x light treatment combination, except in shaded B. papyrifera. The light compensation point (Gamma) decreased in response to shade in all species, and in response to elevated CO(2) in A. rubrum and Q. rubra. Acer rubrum had the greatest increases in apparent quantum yield (phi) in response to shade and elevated CO(2). To illustrate the effects of shifts in A, Gamma and phi on daily C gain, daily integrated C balance was calculated for individual sun and shade leaves. Ignoring possible stomatal effects, estimated daily (24 h) leaf C balance was 218 to 442% higher in the elevated CO(2) treatments than in the ambient CO(2) treatments in both sun and shade seedlings of Q. rubra and A. rubrum. These results suggest that the ability of species to acclimate photosynthetically to elevated CO(2) may, in part, be related to their ability to adapt to low irradiance. Such a relationship has implications for altered C balance and nitrogen use efficiency of understory seedlings.     

Environmental controls over isoprene emission in deciduous oak canopies

In summer 1992, isoprene emission was measured on intact leaves and branches of Quercus alba (L.) at two heights in a forest canopy. Isoprene emission capacity (measured at 30 degrees C and a photosynthetic photon flux density of 1000 micro mol m(-2) s(-1)) was significantly higher in sun leaves than in shade leaves when expressed on a leaf area basis (51 versus 31 nmol m(-2) s(-1); P < 0.01). Because leaf mass per unit area (LMA, g m(-2)) was higher in sun leaves than in shade leaves, emissions of sun and shade leaves expressed on a dry mass basis did not differ significantly (99 versus 89 micro g C g(DW) (-1) h(-1); P = 0.05). Similar measurements in 1995 were consistent with the 1992 data, but data from leaves in more shaded locations demonstrated that isoprene emission capacity decreased with decreasing growth irradiance, irrespective of units of expression. Isoprene emission capacity in leaves of Q. coccinea Muenchh. and Q. velutina Lam. also declined steeply with canopy depth. Emission capacity, on a dry mass basis, showed no obvious pattern with canopy position in Q. prinus L. There was no difference in the temperature response of sun versus shade leaves of Q. alba, but shade leaves exhibited a greater quantum efficiency and saturated at lower irradiance than sun leaves. Rates of isoprene emission measured on branches of Q. alba were approximately 60% of those measured on individual leaves, as a result of self-shading within branch enclosures. It is recommended that within-canopy variation in isoprene emission capacity be incorporated into regional emission models.     

Characterization of the photosynthetic induction response in a Populus species with stomata barely responding to light changes

The photosynthetic induction response is constrained by stomatal and biochemical limitations. However, leaves in some plants like Populus koreana x trichocarpa cv. Peace (a hybrid clone) may have little stomatal limitation because their stomata barely respond to changes in photon flux density (PFD). We examined the induction responses of leaves of well-watered and dehydrated P. koreana x trichocarpa plants grown in a high-light or a low-light regime. With an increase in PFD from 50 to 500 micromol m(-2) s(-1), steady-state stomatal conductance (g(s)) increased by only 0.25-8.2%, regardless of the initial g(s), but steady-state assimilation rate (A) increased by 550-1810%. Photosynthetic induction times required to reach 50% (IT50) and 90% (IT90) of A at high PFD were 60-90 s and 210-360 s, respectively. Examination of the dynamic relationships between A and g(s), and between A and intercellular CO2 concentration, indicated that the induction limitation was imposed completely by the biochemical components within 30-40 s after the PFD increase. Values of IT50 and IT90 were significantly higher in low-light leaves than in high-light leaves, whereas the induction state at 60 s and the induction efficiency at 60 and 120 s after the increase in PFD were lower in low-light leaves than in high-light leaves. Dehydration reduced leaf water potential (psi) significantly, resulting in a significantly decreased initial g(s). Leaf water potential had no significant effects on induction time in high-light leaves, but a low psi significantly reduced the induction time in low-light leaves. We conclude that the photosynthetic induction response was limited almost completely by biochemical components because the stomata barely responded to light changes. The biochemical limitation appeared to be higher in low-light leaves than in high-light leaves. Mild water stress may have reduced steady-state A and g(s), but it had little effect on the photosynthetic induction response in high-light leaves.     

Interactive responses of Quercus suber L. seedlings to light and mild water stress: effects on morphology and gas exchange traits

? The combined effect of water stress and light on seedlings of forest species is a key factor to determine the best silvicultural and afforestation practices in the Mediterranean area.

? The aims of this work was (1) to determine the optimal light level for the early development of cork oak seedlings under mild water stress and (2) to test if the combined effect of water stress and light followed the trade-off, the facilitation or the orthogonal hypothesis.

? Shade reduced instantaneous photosynthetic rates and water use efficiency in cork oak. However, seedlings grown under moderate shade (15% of full sunlight) were capable to accumulate similar amount of biomass than those grown under more illuminated environments by increasing their specific leaf area. Absolute differences in net photosynthesis between light treatments were higher in well watered than in water stressed seedlings. However, the impact of both factors on overall growth was orthogonal.

? We concluded that cork oak development is impaired under deep shade (5% of full sunlight) but it can be optimal under moderate shade (15% of full sunlight) even under moderate water stress. Implications of these patterns on regeneration, cultivation and afforestation of cork oak are discussed.

     

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.     

Seasonal changes in the xanthophyll cycle and antioxidants in sun-exposed and shaded parts of the crown of Cryptomeria japonica in relation to rhodoxanthin accumulation during cold acclimation

Xanthophyll rhodoxanthin, which is present in sun-exposed needles of certain gymnosperms in winter, may have a photoprotective role during long-term cold acclimation. To examine how cold acclimation processes vary within tree crowns and to examine putative correlations between xanthophyll cycle pigments (VAZ), rhodoxanthin and the water-water cycle in photoprotection, we monitored seasonal changes in the activities of two key antioxidant enzymes (ascorbate peroxidase (APX) and glutathione reductase (GR)), pigment composition and chlorophyll fluorescence parameters in sun and shade needles of crowns of the gymnosperm Cryptomeria japonica D. Don. Although APX and GR activities in both sun and shade needles were higher in winter than in summer when assayed at 20 degrees C, differences between seasons were less pronounced when enzymatic activities in summer and winter were assayed at 20 and 5 degrees C, respectively. These results suggest that increases in the potential activity of antioxidant enzymes in winter is an adaptation that helps counterbalance reductions in absolute enzyme activity caused by low temperature, and thus allows the photoprotective capacity of the water-water cycle in C. japonica to be maintained at a roughly constant value throughout the year. In shade needles, the concentration of VAZ increased in winter, but no rhodoxanthin accumulated. Photosynthetic activity was maintained in winter. In sun needles, however, the electron transport rate (ETR) and photochemical quenching (q(P)) decreased to their lowest values in December, just before the accumulation of rhodoxanthin, which coincided with the highest amount of VAZ. Changes in rhodoxanthin concentration mirrored changes in VAZ concentration from January to March. Winter values of ETR and q(P) were comparable with summer values after accumulation of rhodoxanthin, indicating that rhodoxanthin may play a more important role than the VAZ cycle in protecting the photosynthetic apparatus from photodamage in winter. Photosynthetic activity may be modulated, as a result of the interception of light by rhodoxanthin, to match the extent to which absorbed light energy can be utilized in winter when the VAZ cycle is unable to operate effectively because of low temperatures.     

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.     

Hyperhydricity in apple: ultrastructural and physiological aspects

We studied the effects of hyperhydricity on subcellular ultrastructure and physiology of leaves during in vitro regeneration of apple plants. Morphological, anatomical and ultrastructural differences between healthy leaf tissues obtained from greenhouse-grown plants and healthy and hyperhydric leaves obtained from shoots raised from nodal shoot explants in a bioreactor were investigated by electron microscopy and confocal laser scanning microscopy. Compared with healthy leaves, hyperhydric leaves showed abnormal, often discontinuous development of the epidermis and cuticle. Stomata were malformed. The leaf lamina appeared thickened and was characterized by poor differentiation between the palisade and spongy mesophyll tissue. Hyperhydric leaves had a significantly lower chloroplast number per cell and chloroplasts showed reduced thylakoid stacking compared with healthy leaves. Hyperhydricity resulted in a general decrease in concentrations of reduced and oxidized pyridine nucleotides, reflecting a reduction in metabolic activity. The activities of antioxidant enzymes, such as superoxide dismutase, catalase, ascorbate peroxidase and glutathione reductase were higher in hyperhydric leaves than in healthy leaves, indicating that hyperhydricity was associated with oxidative stress. Chlorophyll fluorescence measurements provided evidence of oxidative damage to the photosynthetic machinery in hyperhydric leaves: photochemical efficiency of photosystem II, effective quantum efficiency and photochemical quenching were all lower in hyperhydric leaves compared with healthy leaves.     

Effects of light availability on leaf gas exchange and expansion in lychee (Litchi chinensis)

Effects of photosynthetic photon flux density (PPFD) on leaf gas exchange of lychee (Litchi chinensis Sonn.) were studied in field-grown "Kwai May Pink" and "Salathiel" orchard trees and young potted "Kwai May Pink" plants during summer in subtropical Queensland (27 degrees S). Variations in PPFD were achieved by shading the trees or plants 1 h before measurement at 0800 h. In a second experiment, potted seedlings of "Kwai May Pink" were grown in a heated greenhouse in 20% of full sun (equivalent to maximum noon PPFD of 200 micromol m(-2)xs(-1)) and their growth over three flush cycles was compared with seedlings grown in full sun (1080 micromol m(-2)xs(-1)). Young potted plants of "Kwai May Pink" were also grown outdoors in artificial shade that provided 20, 40, 70 or 100% of full sun (equivalent to maximum PPFDs of 500, 900, 1400 and 2000 micromol m(-2)xs(-1)) and measured for shoot extension and leaf area development over one flush cycle. Net CO2 assimilation increased asymptotically in response to increasing PPFD in both orchard trees and young potted plants. Maximum rates of CO2 assimilation (11.9 +/- 0.5 versus 6.3 +/- 0.2 micromol CO2 m(-2) s(-1)), dark respiration (1.7 +/- 0.3 versus 0.6 +/- 0.2 micromol CO2 m(-2) s(-1)), quantum yield (0.042 +/- 0.005 versus 0.027 +/- 0.003 mol CO2 mol(-1)) and light saturation point (1155 versus 959 micromol m(-2) s(-1)) were higher in orchard trees than in young potted plants. In potted seedlings grown in a heated greenhouse, shoots and leaves exposed to full sun expanded in a sigmoidal pattern to 69 +/- 12 mm and 497 +/- 105 cm(2) for each flush, compared with 27 +/- 7 mm and 189 +/- 88 cm(2) in shaded seedlings. Shaded seedlings were smaller and had higher shoot:root ratios (3.7 versus 3.1) than seedlings grown in full sun. In the potted plants grown outdoors in 20, 40, 70 or 100% of full sun, final leaf area per shoot was 44 +/- 1, 143 +/- 3, 251 +/- 7 and 362 +/- 8 cm(2), respectively. Shoots were also shorter in plants grown in shade than in plants grown in full sun (66 +/- 5 mm versus 101 +/- 2 mm). Photosynthesis in individual leaves of lychee appeared to be saturated at about half full sun, whereas maximum leaf expansion occurred at higher PPFDs. We conclude that lychee plants can persist as seedlings on the forest floor, but require high PPFDs for optimum growth.