Acclimation of leaves to contrasting irradiance in juvenile trees differing in shade tolerance

Leaves developing in different irradiances undergo structural and functional acclimation, although the extent of trait plasticity is species specific. We tested the hypothesis that irradiance-induced plasticity of photosynthetic and anatomical traits is lower in highly shade-tolerant species than in moderately shade-tolerant species. Seedlings of two evergreen conifers, shade-tolerant Abies alba Mill. and moderately shade-tolerant Picea abies Karst., and two deciduous angiosperm species, highly shade-tolerant Fagus sylvatica L. and moderately shade-tolerant Acer pseudoplatanus L., were grown in deep shade (LL, 5% of full irradiance) or in full solar irradiance (HL) during 2003 and 2004. Steady state responses of quantum yield of PSII (Phi(PSII)), apparent electron transport rate (ETR), nonphotochemical quenching (NPQ) and photochemical quenching (qP) were generally modified by the light environment, with slower declines in Phi(PSII) and qP and greater maximal ETR and NPQ values in HL plants in at least one season; however, no link between quantitative measures of plasticity of these traits and shade tolerance was found. Plasticity of nine anatomical traits (including palisade cell length, which was reduced in LL) showed no relationship with shade tolerance, but was less in conifers than in deciduous trees, suggesting that leaf life span may be a significant correlate of plasticity. When LL-acclimated plants were exposed to HL conditions, the degree and duration of photoinhibition (measured as a decline in maximum quantum yield) was greatest in F. sylvatica, much lower in P. abies and A. alba, and lowest in A. pseudoplatanus. Thus, as with the other traits studied, vulnerability to photoinhibition showed no relationship with shade tolerance.     

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.     

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.     

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 nursery shading on seedling quality and post-planting performance in two Mediterranean species with contrasting shade tolerance

In Mediterranean climates, seedlings are frequently shaded in the nursery to avoid heat damage and save water. However, the impact of this shading on the seedling quality and transplanting performance of Mediterranean species is not well known. We studied the effect of nursery shading on pre-planting features and post-planting performance of two Mediterranean tree species: the shade-intolerant pioneer Pinus halepensis and the shade-tolerant late-successional Quercus ilex. We grew one-year-old seedlings of both species under 100, 40 and 5% full sunlight. Shade had a low impact on the morphology and physiology of Q. ilex seedlings. In pines, only the deep shade treatment produced low quality seedlings with poor root development. In both species, transference to high light at planting in autumn did not impose any additional stress than that caused by frosts, but initial root growth was impaired in the two shaded treatments in pine. Post-planting growth and survival of oak seedlings showed no difference between treatments. Pine seedlings grown in deep shade showed higher mortality and lower growth after planting than those grown in full sun and intermediate light treatments, while intermediate light only reduced growth. For the nursery culture of Q. ilex seedlings, we advise using low light levels during summer to save water without impairing field performance. In P. halepensis, seedlings should be cultured under full sunlight conditions to maximize post-planting growth, but they can be cultured under intermediate light without impairing survival.     

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.     

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.     

Functional traits and plasticity in response to light in seedlings of four Iberian forest tree species

We investigated the differential roles of physiological and morphological features on seedling survivorship along an experimental irradiance gradient in four dominant species of cool temperate-Mediterranean forests (Quercus robur L., Quercus pyrenaica Willd., Pinus sylvestris L. and Pinus pinaster Ait.). The lowest photochemical efficiency (F(v)/F(m) in dark-adapted leaves) was reached in deep shade (1% of full sunlight) in all species except Q. robur, which had the lowest photochemical efficiency in both deep shade and 100% of full sunlight. Species differed significantly in their survival in 1% of full sunlight but exhibited similar survivorship in 6, 20 and 100% of full sunlight. Shade-tolerant oaks had lower leaf area ratios, shoot to root ratios, foliage allocation ratios and higher rates of allocation to structural biomass (stem plus thick roots) than shade-intolerant pines. Overall phenotypic plasticity for each species, estimated as the difference between the minimum and the maximum mean values of the ecophysiological variables studied at the various irradiances divided by the maximum mean value of those variables, was inversely correlated with shade tolerance. Observed morphology, allocation and plasticity conformed to a conservative resource-use strategy, although observed differences in specific leaf area, which was higher in shade-tolerant species, supported a carbon gain maximization strategy. Lack of a congruent suite of traits underlying shade tolerance in the studied species provides evidence of adaptation to multiple selective forces. Although the study was based on only four species, the importance of ecophysiological variables as determinants of interspecific differences in survival in limiting light was demonstrated.     

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.     

Acclimation of Pacific yew (Taxus brevifolia) foliage to sun and shade

The success in clinical trials of the anti-cancer drug, Taxol(R), obtained from the bark of Pacific yew (Taxus brevifolia Nutt.), has raised interest in cultivation and regeneration of this little-known species. Pacific yew is shade-tolerant and it is not known whether the foliage can tolerate the high solar irradiances found on an open forest regeneration site or a nursery. Acclimation of Pacific yew to sun and shade was studied by comparing foliar physiology and morphology of male and female trees growing in full sun or shade. Interspecific foliar acclimation to sun was studied by comparing sun-grown English yew (Taxus baccata L.) with Pacific yew. No sex-specific acclimation was found in foliar physiology or morphology in either species. Sun-grown foliage of Pacific yew and English yew differed with respect to light harvesting, transpiration, stomatal conductance, leaf structure, stomatal distribution and foliar N concentrations and contents. Chlorophyll a fluorescence measurements indicated that shade-grown foliage of Pacific yew had larger and more efficient light harvesting systems than sun-grown foliage. Rates of CO(2) uptake and transpiration were similar in sun- and shade-grown foliage indicating acclimation of photosynthesis to the growth irradiance. Specific leaf area was significantly higher in shade-grown foliage of Pacific yew than in sun-grown foliage and was diagnostic of the light environment in which the foliage grew. Foliar N concentrations were not significantly different between sun- and shade-grown leaves of Pacific yew but sun-grown foliage had a higher N content. Physiological and morphological adjustments of Pacific yew foliage conferred tolerance to both high light and shade, enabling the trees to survive in a variety of light environments and indicating that Pacific yew is suited to nursery cultivation and regeneration of open sites.     

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.     

Characteristics of the Light Response in Seedlings and Saplings of Two Mid-Successional Species,Ash and Kalopanax,During the Early Stage of Regeneration in a Mature Forest

Abstract

Growth characteristics of ash (Fraxinus mandshuricavar. japonicd)and kalopanax (Kalopanax pictus), two mid-succcs-sional species, were monitored in relation to seasonal light in a deciduous broadleaf forest. During the growing, relative light intensity of the gap was 4 to 6 times higher than that under the closed canopy. Seedlings could be found on the mixed hardwood forest floor where relative light intensity was around greater than 7% of full sunlight. When saplings gap openings which had relative light intensities above 20%, they quickly developed lateral branches and increased their foliage volume. The light-photosynthesis curve of seedlings of both species showed a curve typical of the “shade leaf” type even though they grew under sunlight. In contrast, saplings of both species showed the “sun leaf” type of light-photosynthesis curve. This seedling to sapling shift from shade to sun adaptation was also found in other foliage characteristics. This study shows that species can efficiently adjust their leaf and branch characteristics to changing light environments from shade to openings in a forest.     

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.     

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.     

Water relations of seedlings of three Quercus species: variations across and within species grown in contrasting light and water regimes

We compared seedling water relations of three Mediterranean Quercus species (the evergreen shrub Q. coccifera L., the evergreen tree Q. ilex L. subsp. ballota (Desf.) Samp. and the deciduous or marcescent tree Q. faginea L.). We also explored seedling potential for acclimation to contrasting growing conditions. In March, 1-year-old seedlings of the three species were planted in pots and grown outdoors in a factorial combination of two irrigation regimes (daily (HW) and alternate day watering (LW)) and two irradiances (43 and 100% of full sunlight). At the end of July, predawn and midday water potentials (Psi(pd), Psi(md)) were measured, and pressure-volume (P-V) curves were obtained for mature current-year shoots. Species exhibited similar Psi(pd) and Psi(md) values, but differed in leaf morphology and water relations. The evergreens possessed larger leaf mass per area (LMA) and were able to maintain positive turgor pressure at lower water potentials than the deciduous species because of their lower osmotic potential at full turgor. However, the three species had similar relative water contents at the turgor loss point because Q. faginea compensated for its higher osmotic potential with greater cell wall elasticity. Values of Psi(pd) had a mean of -1.12 MPa in LW and -0.63 MPa in HW, and Psi(md) had a mean of -1.13 MPa in full sunlight and -1.64 MPa in shade, where seedlings exhibited lower LMA. However, the P-V curve traits were unaffected by the treatments. Our results suggest that Q. faginea seedlings combine the water-use characteristics of mesic deciduous oak and the drought-tolerance of xeric evergreen oak. The ability of Q. coccifera to colonize drier sites than Q. ilex was not a result of higher drought tolerance, but rather may be associated with other dehydration postponement mechanisms including drought-induced leaf shedding. The lack of treatment effects may reflect a relatively low contrast between treatment regimes, or a low inherent responsiveness of these traits in the study species, or both.     

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.     

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.     

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.     

Photosynthetic responses to understory shade and elevated carbon dioxide concentration in four northern hardwood tree species

Seedling responses to elevated atmospheric CO(2) concentration ([CO(2)]) and solar irradiance were measured over two growing seasons in shade-tolerant Acer saccharum Marsh. and Fagus grandifolia J.F. Ehrh. and shade-intolerant Prunus serotina, a J.F. Ehrh. and Betula papyrifera Marsh. Seedlings were exposed to a factorial combination of [CO2] (ambient and elevated (658 micromol mol-1)) and understory shade (deep and moderate) in open-top chambers placed in a forest understory. The elevated [CO(2)] treatment increased mean light-saturated net photosynthetic rate by 63% in the shade-tolerant species and 67% in the shade-intolerant species. However, when measured at the elevated [CO(2)], long-term enhancement of photosynthesis was 10% lower than the instantaneous enhancement seen in ambient-[CO(2)]-grown plants (P < 0.021). Overall, growth light environment affected long-term photosynthetic enhancement by elevated [CO(2)]: as the growth irradiance increased, proportional enhancement due to elevated [CO(2)] decreased from 97% for plants grown in deep shade to 47% for plants grown in moderate shade. Results suggest that in N-limited northern temperate forests, trees grown in deep shade may display greater photosynthetic gains from a CO(2)-enriched atmosphere than trees growing in more moderate shade, because of greater downregulation in the latter environment. If photosynthetic gains by deep-shade-grown plants in response to elevated [CO(2)] translate into improved growth and survival of shade-intolerant species, it could alter the future composition and dynamics of successional forest communities.     

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.