Comparison of the physiology, morphology, and leaf demography of tropical saplings with different crown shapes

Branch architecture, leaf photosynthetic traits, and leaf demography were investigated in saplings of two woody species, Homolanthus caloneurus and Macaranga rostulata, co-occurring in the understory of a tropical mountain forest. M. rostulata saplings have cylindrical crowns, whereas H. caloneurus saplings have flat crowns. Saplings of the two species were found not to differ in area-based photosynthetic traits and in average light conditions in the understory of the studied site, but they do differ in internode length, leaf emergence rate, leaf lifespan, and total leaf area. Displayed leaf area of H. caloneurus saplings, which have the more rapid leaf emergence, was smaller than that of M. rostulata saplings, which have a longer leaf lifespan and larger total leaf area, although M. rostulata saplings showed a higher degree of leaf overlap. Short leaf lifespan and consequent small total leaf area would be linked to leaf overlap avoidance in the densely packed flat H. caloneurus crown. In contrast, M. rostulata saplings maintained a large total leaf area by producing leaves with a long leaf lifespan. In these understory saplings with a different crown architecture, we observed two contrasting adaptation strategies to shade which are achieved by adjusting a suite of morphological and leaf demographic characters. Each understory species has a suite of morphological traits and leaf demography specific to its architecture, thus attaining leaf overlap avoidance or large total leaf area.     

Carbon Translocation as Affected by Shade in Saplings of Shade Tolerant and Intolerant Species

Carbon translocation was affected by shade in different tropical tree species differing in successional status and degree of shade tolerance. Plants of the early-successional shade-intolerant species Cecropia pachystachya and Schizolobium parahyba and of the late-successional shade-tolerant species Myroxylon peruiferum and Hymenaea courbaril were grown under full sun (FS) and natural shade treatments (NS) and assessed for [¹⁴C]-sucrose translocation. Most of the ¹⁴C was retained in the fed leaf after a 24 h translocation period. Under FS, the growing apical part of the plant was the most intense sink for most species. Shade affected growth and sink intensity differently in early and late successional species. Growth was more markedly affected in the early species. Whereas these continued to invest carbon into the growing apical part of the plant under shade conditions, the late successional species invested relatively more into other sinks. This revised version was published online in July 2006 with corrections to the Cover Date.     

Photosynthesis in an Australian rainforest tree,Argyrodendron peralatum,during the rapid development and relief of water deficits in the dry season

Summary Rates of apparent photosynthesis were measured in situ at five positions between the upper crown and a lower branch of a 34 m tall Argyrodendron peralatum (F.M. Bailey) H.L. Edlin ex I.H. Boas tree, and on an understorey sapling of the same species growing in a northern Australian rainforest. At the end of the dry season, rapid reductions in photosynthetic rates occurred in the upper crown within three days after a rain event, but changes in the lower crown and the sapling were less marked. Complete recovery of photosynthesis followed a second rain event. At high photon flux densities, stomatal conductance to water vapour decreased in a curvilinear fashion as the vapour pressure difference between leaf and air increased. Apparent photosynthesis was linearly related to stomatal conductance on the first clear day after each rain event, but there was no relationship between these parameters at the end of a brief natural drying cycle. Under conditions of adequate water supply, stomatal conductances of both upper crown and understorey leaves increased linearly with increasing photon flux density up to about 300 mol m^-2 s^-1. During water deficits, stomatal conductances in leaves from the understorey increased much more rapidly at very low photon flux densities than did conductances in leaves from the upper canopy.     

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

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

Mid-successional stand dynamics in a cool-temperate conifer-hardwood forest in northern Japan

Stand dynamics was studied over 13 years in a cool-temperate conifer-hardwood forest, northern Japan. A total 30 hardwood species and one conifer, Abies sachalinensis, larger than 1.5 cm DBH were recorded. The total stand density was 1677 trees ha⁻¹ at the beginning, decreasing to 1184 trees ha⁻¹ (30% reduction) over the study period, but the total stand basal area was almost unchanged (about 49 m² ha⁻¹). This large reduction in total density was mainly due to the death of saplings and infrequent recruitment. Number of recruits gradually decreased with time, while that of dead trees was constant. Cause of death of small trees was mainly due to suppression by tall trees. Skewness of the DBH frequency distribution varied among the species. A less skewed frequency distribution (i.e., few number of saplings) was shown by shade-intolerant species such as Populus maximowiczii and Betula maximowicziana, and a more skewed frequency distribution (i.e., large number of saplings) by shade-tolerant species such as Acer mono and Tilia japonica. DBH frequency distribution changed to less skewed patterns with reduction of density in most species during the census period. Rank of shade tolerance positively correlated with tree density and skewness, and negatively correlated with mean DBH. Skewness also positively correlated with recruitment rates. Furthermore, rank of shade tolerance positively correlated with seed size. These results suggest that shade-intolerant species regenerated immediately after disturbances by wide dispersal of small seeds, but their recruitment was interrupted after that. By contrast, shade-tolerant species were able to recruit even after the ceasation of recruitment of shade-intolerant species, but suffered severe mortality due to the increasing shading with the progress of stand development. This study suggests that the stand is still developing, with changes in species composition and size structure, and that species differences in shade tolerance and seed size are important for the stand structural changes.     

Multiple canopy opening effects on recruited saplings in a typhoon-disturbed tropical rainforest,Taiwan

To investigate the influence of multiple canopy openings on the composition and diversity of recruited saplings in a forest frequently disturbed by typhoons. We conducted tree-by-tree censuses (diameter at breast height ≥ 1 cm) and mapped gaps (canopy height < 5 m) in 1993, 2000, 2008, and 2013 in a tropical mountain zonal foothill evergreen broad-leaved forest in Nanjenshan Nature Reserve, Taiwan. We analyzed the composition and diversity of recruited saplings within a 2.1 ha plot (840 sampling quadrats (5 m × 5 m)) with variable numbers of canopy openings recorded during the study period. Composition of recruited saplings was dissimilar between quadrats that stayed opened and those that stayed closed throughout the study period (pairwise similarity estimates C₀₂ = 0.52, 95% CI = 0.38–0.66). The quadrats under closed canopy had high diversity when weighting rare species (species richness), whereas quadrats with one or two gap opening records during the past 20 years had high diversity when weighting the abundance of species. Although canopy openings provided establishment conditions for saplings of some shade-intolerant species, due to the nature of small gap size, such habitats do not favor the dominance of shade-intolerant species. Even in a frequently disturbed forest, species composition and richness of recruited saplings were mainly contributed by shade-tolerant species. Although multiple canopy openings facilitated the establishment of shade-intolerant species, species diversity in the study forests is possibly mainly mediated by coexistence mechanisms of those shade-tolerant species rather than light-gap-related species strategies.     

Dwarf bamboos affect the regeneration of zoochorous trees by providing habitats to acorn-feeding rodents

The effects of dwarf bamboos (Sasa spp.) on the regeneration of trees in a natural hardwood forest were studied by analysing the spatial dispersion of seedlings and saplings of anemochores (Acer palmatum var. matsumurae, Fraxinus lanuginosa, and Carpinus laxiflora) and zoochores (Quercus mongolica var. grosseserrata and Q. serrata). Relative photosynthetic photon flux density at 10 cm above ground was significantly correlated with the coverage of dwarf bamboos (r=0.661, P<0.001). Seedlings were abundant and were randomly distributed in the anemochores, other than the shade-intolerant species C. laxiflora which was significantly more sparse in sites with dense Sasa than in sites where Sasa was rare. Distribution of saplings was also random in the shadetolerant anemochores A. palmatum var. matsumurae and F. lanuginosa but aggregated in sites with sparse Sasa in the shade-intolerant anemochore C. laxiflora. In contrast to the anemochores, seedlings of zoochores were very few and were distributed in sites with sparse Sasa. Saplings were also aggregated and negatively correlated with Sasa cover in the shade-intolerant species Q. serrata and the tolerant species Q. mongolica var. grosseserrata. The acorns put on the forest floor in a site with dense Sasa were quickly removed by small rodents such as Apodemus speciosus and A. argenteus. Trap census of rodents revealed that those mammals prefer the dense Sasa habitat to the sparse Sasa habitat. This suggests that the dwarf bamboos strongly affect the regeneration of zoochorous trees not only by shading the seedlings but also by providing habitats to acorn-feeding small mammals.     

Differences in sapling architecture betweenFagus crenata andFagus japonica

The ecological significance of architectural patterns for saplings ofFagus crenata andFagus japonica co-occurring in a secondary oak forest were evaluated by comparing the size and shape of leaves, trunks and crowns.Fagus japonica saplings were different fromF. crenata saplings in some architectural properties: (i) the leaf area and specific leaf area were larger; (ii) the ratio of sapling height to trunk length was lower, indicating greater leaning of the trunk; and (iii) the projection area of the crown was larger and the leaf area index lower indicating less mutual shading of leaves. These architectural features indicated thatF. japonica saplings were more shade tolerant thanF. crenata andF. crenata saplings were superior toF. japonica for growth in height and could, therefore, utilize sunlight in the upper layer. An erect trunk inF. crenata and a leaning trunk inF. japonica may be important characteristics associated with the regenerations patterns of each species; regeneration from seedlings under canopy gaps in the former and vegetative regeneration by sprouting in the latter.     

Photosynthesis and leaf morphology ofLiquidambar styraciflua L. under variable urban radiant-energy conditions

Diminished sunlight, characteristic of urban canyons, has been suggested as being potentially limiting to plant growth. This study investigated the response of sweetgum (Liquidambar styraciflua L.) to variable irradiance in a range of urban locations. Diurnal photosynthesis was measured in situ on mature trees, comparing an open site at an urban park with an urban canyon that received 4 h of midday sun in midsummer. Photosynthesis for trees growing in the canyon was lower both during shaded and sunlit periods compared with trees at the park. Photosynthesis of detached shoots in a growth chamber was greater in canyon than park foliage at low irradiance, indicating possible photosynthetic shade acclimation analogous to tree species growing in the forest understorey. Shoot and trunk growth and morphological characteristics were measured onL. styraciflua growing along boulevards at 15 additional urban sites and related to seasonal interception of solar radiation. Angular elevation and orientation of buildings and trees that defined the horizon topography at each site were used in modeling the potential irradiance of global shortwave radiation. Seasonal irradiance among sites ranged from 21% in the urban core to nearly 95% in outlying residential districts of that potentially received under an unobstructed horizon. Shade acclimation was confirmed by differences in leaf morphology, as foliage became flatter, thinner, and more horizontally oriented at sites with lower irradiance. Photosynthetic and morphological acclimation to shade did not compensate for lower available radiant energy as both shoot and trunk growth decreased at sites of lower irradiance. Unlike the forest understorey, the static light environment of urban canyons may subject shade-intolerant species such asL. styraciflua to chronic, low-radiant-energy stress.     

The functional morphology of juvenile plants tolerant of strong summer drought in shaded forest understories in southern Spain

It has been hypothesized that plants cannot tolerate combined shade and drought, as a result of morphological trade-offs. However, numerous plant species are reportedly widespread in shaded forest understories that face drought, whether seasonal or occasional. We studied juveniles of six plant species that cope with strong summer drought in the understoreys of mixed Quercus forests in southern Spain: the tall-shrubs Phillyrea latifolia and Viburnum tinus, the perennial herb Rubia peregrina, the small shrub Ruscus aculeatus, and climbers Hedera helix and Smilax aspera. All of these species persist in evergreen shade (c. 3% daylight). Two other species were studied as comparators, Ruscus hypoglossum, less tolerant of drought, and Ceratonia siliqua, less tolerant of shade. Morphological and chemical variables relevant to shade and drought tolerance were measured for juveniles in a range of sizes, and also for the leaves of mature plants. The species converge in features that confer tolerance of shade plus drought by reducing demand for resources. Demand for water is reduced through a moderate to high below-ground mass fraction and low to moderate specific leaf area (respectively 0.22–0.52 and 112–172 cm² g^–1 at 1.00 g total dry mass). Demand for both irradiance and water is reduced through a low to moderate foliar nitrogen concentration and long-lived, physically protected leaves (2 yr). The species also converge in features that confer tolerance of either low irradiance or drought through specialized capture of resource, without precluding the other tolerance. These features include deep roots relative to shoot size, moderately higher specific leaf area in shade (1.2–2.0 × that in sun) and higher chlorophyll:nitrogen ratio in shade. Foliar chlorophyll per unit mass was higher in shade, but chlorophyll was not necessarily synthesized in greater amounts; rather, it was higher apparently due to shade effects on structural features linked with specific leaf area. In contrast, N per unit mass was higher in sun leaves independently of specific leaf area. Despite these convergences, the species diverge considerably in their root mass allocation and architecture, leaf saturated water content, density of stomata and guard cell size. No single narrowly defined functional type is needed for tolerance of shade plus drought.     

Photosynthetic induction in leaves of co-occurring <Emphasis Type="Italic">Fagus lucida</Emphasis> and <Emphasis Type="Italic">Castanopsis lamontii</Emphasis> saplings grown in contrasting light environments

Photosynthetic induction times and photoinhibition in relation to simulated sunflecks (sudden increase of irradiance from 20 to 1,500 μmol m⁻² s⁻¹) were examined in leaves of co-occurring Fagus lucida (a deciduous tree) and Castanopsis lamontii (an evergreen tree) saplings grown either in a beech forest understory or in an adjacent open site during a late rainy season. Two hypotheses were tested: (1) understory leaves would display faster photosynthetic induction times and greater photoinhibition than open-grown leaves; and (2) evergreen species would have slower photosynthetic induction times and lighter photoinhibition than deciduous species. Times to reach 90% of maximal CO₂ assimilation rate (t _90%A ) and stomatal conductance did not differ between species, but showed faster by 3–5 min in open-grown leaves than understory leaves due to higher initial stomatal conductance (g _{s initial}) and induction state 1 min into simulated sunflecks (IS_1min) in the former. Our analysis across the published data on photosynthetic induction of 48 broad-leaved woody species again revealed the negative correlations between t _90%A and either g _{s initial} or IS_1min, and the similarity of t _90%A and between evergreen and deciduous species. Measurements of maximum PSII photochemical efficiency (F _v/F _m) indicated that photoinhibition occurred in saplings in any of the growth habitats during sunfleck-induced photosynthetic induction. Despite no interspecific differences in the degree of photoinhibition, understory leaves of both species suffered heavier photoinhibition than open-grown leaves, as indicated by a stronger decrease of F _v/F _m in the former. Dynamic changes in the quantum yields of PSII photochemistry and ΔpH- and xanthophyll-regulated thermal dissipation and adjustments in the partitioning of electron flow between assimilative and non-assimilative processes were functional to resist photoinhibition. However, such photoinhibition, together with stomatal and biochemical limitations, would decrease carbon gain during simulated sunflecks, particularly in understory leaves.     

Shade adaptation and shade tolerance in saplings of three Acer species from eastern North America

Summary Saplings of three, co-occurring maple species in a mature maple-beech forest differed in a suite of structural and physiological characters that separated the canopy species, Acer, saccharum, from the two subcanopy species, A. pensylvanicum and A. spicatum. Acer saccharum had both more dense wood and tougher and heavier but thinner leaves than the subcanopy species. Acer pensylvanicum had the largest, lightest leaves with high stomatal density and its canopy architecture was the most effective in terms of leaf display for light interception. Acer spicatum had weaker wood similar to that of A. pensylvanicum but also small, soft and relatively poorly displayed leaves. Both subcanopy species maintained marginally higher average rates of photosynthesis over the growing season in the understory environment. We consider juvenile A. saccharum only shade-tolerant, capable of persisting through long periods in the closed canopy until a gap occurs but not specifically adapted to the understory environment. Juvenile A. sacchrum appears to be constrained functionally by the requirements set by the canopy environment that adults will occupy. Characters such as high wood density are already expressed in the understory sapling; this investment in denser wood slows the growth of saplings, but is necessary for structural reasons in the adult. Juvenile A. saccaharum have morphological and photosynthetic characters better suited to gas exchange and extension growth under the increased photon flux densities in large forest gaps, characteristics that will also be advantageous in the sunlit canopy environment of adults.Both subcanopy maples appear to be more truly shade-adapted, although in somewhat different ways. Acer pensylvanicum has characteristics that enhance the potential for capture and utilization of sunflecks and is able to sustain higher growth rates than A. saccharum in the shaded subcanopy environment. Acer spicatum shares some shade-adapted features with A. pensylvanicum, and its habit of lateral spread through stem layering may confer an additional advantage in foraging for small light gaps.     

Allometry, root/shoot ratio and root architecture in understory saplings of deciduous dicotyledonous trees in central Japan

Plant allometry that is related to plant architecture and biomass allocation strongly influences a plants ability to grow in shaded forest understory. Some allometric traits can change with plant size. The present study compared crown and trunk allometries, root/shoot biomass allometry, and root architecture among understory saplings (0.25--5m height, except for two trees > 5 < 7 m) of seven deciduous dicotyledonous species in central Japan. Associations of the crown and trunk allometries with several plant morphological attributes were analyzed. Branch morphology (plagiotropyvs orthotropy) and life size were correlated with sapling crown and trunk allometries. Both large leaves and orthotropic branches were associated with a narrow small crown and slender trunk. The root/shoot ratio decreased rapidly with increasing plant height for saplings shorter than about 1.5 m. Less shade-tolerant species tended to have smaller root/shoot ratios for saplings taller than 1.5 m. With an increase in plant height, the branch/trunk biomass ratio decreased for saplings with plagiotropic branches but increased for saplings with orthotropic branches. Four subcanopy species (Acer distylum, Carpinus cordata, Fraxinus lanuginosa and Acanthopanax sciadophylloides) had superficial root systems; a common understory species (Sapium japonica) had a deep tap root system; and a canopy species (Magnolia obovata) and a subcanopy species (Acer tenuifolium) had heart root systems of intermediate depth. The root depth was not related to shade tolerance. Among species of the same height, the difference in fine root length can be 30-fold.     

Determination of the quantum efficiency of photosystem II and of non-photochemical quenching of chlorophyll fluorescence in the field

A newly developed portable chlorophyll fluorometer in combination with a special leaf clip holder was used for assessing photosynthetic activity of attached sun leaves of Fagus sylvatica and Cucurbita pepo under field conditions. During diurnal time courses, fluorescence yield, photosynthetic photon flux density (PPFD) incident on the leaf plane, and leaf temperature were measured and quantum efficiency of photosystem II (PS II), apparent relative electron transport rates, and non-photochemical fluorescence quenching (NPQ) calculated. In both species, quantum efficiency followed closely the incident PPFD and no hysteresis could be observed during the day. Apparent electron transport rate showed light saturation above a PPFD of 700 mol m^–2 s^–1 in F. sylvatica, while in C. pepo no saturation was visible up to 1400 mol m^–2 s^–1. NPQ was closely correlated to excessive PPFD calculated from the PS II quantum yield. Maximal NPQ observed was 3.3 Although the beech leaf was exposed for a considerable time to PPFD values of 1400–1500 mol m^–2 s^–1 and leaf temperatures between 30 and 35°C, no obvious signs for sustained photodamage could be observed. The data demonstrate the potential of chlorophyll fluorescence measurements to analyse photosynthetic performance under field conditions with minimal disturbance of the plant. Potential error sources due to the geometry of the leaf clip holder used are discussed.Dedicated to Prof. Dr. F.-C. Czygan on the occasion of his 60th birthday     

The occurrence of flowering and fruiting on individual trees over 3 years and their effects on subsequent crown condition

Summary The level of fruiting in four forest trees species (Picea sitchensis, P. abies, Pinus sylvestris and Fagus sylvatica) was monitored in Great Britain over the period 1989–1991. In addition, assessments of crown transparency were available for many of the trees for 1987 and 1988. The monitoring period encompassed severe summer droughts in 1989 and 1990, with wetter conditions in 1991. Variations in the level of fruiting in spruce and beech (Fagus sylvatica L.) were seen, with a marked peak in 1990. No pattern was apparent in Scots pine (Pinus sylvestris L.). Coning, which was greater in trees with the least transparent crowns, had no discernible effect on the crown transparency of the conifers. Cupule production in beech was greatest in trees with the most transparent crowns, and trees with high numbers of cupules in 1990 tended to have greater crown dieback recorded in 1991.     

Stand dynamics of the shade-tolerant conifers Podocarpus nubigena and Saxegothaea conspicua in Chilean temperate rain forest

Abstract. Population structures were used to infer regeneration patterns of the conifers Podocarpus nubigena and Saxegothaea conspicua and associated angiosperm tree species in six stands in the temperate rainforests of the coastal ranges of southern Chile. On poorly-drained, nutrient-poor and high altitude sites, where most of the associated species were fairly shade-intolerant and light-crowned, dense all-aged populations and the presence of numerous saplings beneath the canopy suggested continuous regeneration of both P. nubigena and S. conspicua. On more favourable sites, where several of the associated angiosperms were highly shade-tolerant and densecrowned, both conifers, particularly P. nubigena, were less abundant, and their regeneration from seed appeared to be sporadic. S. conspicua's great longevity (≥ 750 yr) and frequent vegetative reproduction are probably crucial to its persistence in competition with shade-tolerant broadleaved species in undisturbed stands on favourable sites. Results of this study, and a review of related literature on southern temperate forests are not consistent with the hypothesis that heavily-shaded, infrequently-disturbed habitats are an evolutionary refuge for conifers. Sites likely to have high leaf area indices and infrequent disturbance are better exploited by shade-tolerant angiosperms.     

Morphological plasticity of regeneration subject to different levels of canopy cover in mixed-species, multiaged forests of the Romanian Carpathians

Morphological plasticity was studied for advanced regeneration trees in different light environments of the mountainous, mixed-species forests in the Carpathian Mountains of Romania. The primary species in these mixtures were very shade tolerant silver fir (Abies alba Mill.) and European beech (Fagus sylvatica L.), and midtolerant Norway spruce (Picea abies (L.) Karst). Seedlings/saplings of these species were selected for measurements in different stands from two different geographical locations. Various morphological traits (specific leaf area, live crown ratio, crown width to length ratio, terminal to lateral ratio, number of internodal shoots, number of shoots in terminal whorl, stem symmetry, stem orientation, stem forking) for each regenerating tree were measured during summers of 2001 and 2002. Percentage of above canopy light and stand basal area measures were used to assess the available growing space for each seedling/sapling. Regression relationships were developed for the different morphological indicators as a function of these two variables. All species adapted their morphology along the gradient in light and basal area. Spruce seemed to be less adapted to low light conditions than both fir and beech. However, no significant differences in terms of shade tolerance were detected using the above indicators. In really dense stand conditions (less than 20% above canopy light and stand basal area above 36 m² ha⁻¹), probability for stem forking in beech increased. In open, all three species adapted their morphology for vigorous growth. Under such conditions, spruce was better adapted than fir.     

Photosynthetic Characteristics,Dark Respiration,and Leaf Mass Per Unit Area in Seedlings of Four Tropical Tree Species Grown Under Three Irradiances

We investigated the effect of growth irradiance (I) on photon-saturated photosynthetic rate (P _max), dark respiration rate (R _D), carboxylation efficiency (CE), and leaf mass per unit area (LMA) in seedlings of the following four tropical tree species with contrasting shade-tolerance. Anthocephalus chinensis (Rubiaceae) and Linociera insignis (Oleaceae) are light-demanding, Barringtonia macrostachya (Lecythidaceae) and Calophyllum polyanthum (Clusiaceae) are shade-tolerant. Their seedlings were pot-planted under shading nets with 8, 25, and 50 % daylight for five months. With increase of I, all species displayed the trends of increases of LMA, photosynthetic saturation irradiance, and chlorophyll-based P _max, and decreases of chlorophyll (Chl) content on both area and mass bases, and mass-based P _max, R _D, and CE. The area-based P _max and CE increased with I for the light-demanders only. Three of the four species significantly increased Chl-based CE with I. This indicated the increase of nitrogen (N) allocation to carboxylation enzyme relative to Chl with I. Compared to the two shade-tolerants, under the same I, the two light-demanders had greater area- and Chl-based P _max, photosynthetic saturation irradiance, lower Chl content per unit area, and greater plasticity in LMA and area- or Chl-based P _max. Our results support the hypothesis that light-demanding species is more plastic in leaf morphology and physiology than shade-tolerant species, and acclimation to I of tropical seedlings is more associated with leaf morphological adjustment relative to physiology. Leaf nitrogen partitioning between photosynthetic enzymes and Chl also play a role in the acclimation to I.     

Diurnal changes in chlorophyll fluorescence and light utilization in Colocasia esculenta leaves grown in marshy waterlogged area

Diurnal cycle of chlorophyll fluorescence parameters was done in Colocasia esculenta L. (swamp taro) grown in marshy land under sun or under shade. The sun leaves maintained higher electron transport rate (ETR) and steady state to initial fluorescence ratio (F_s/F₀) than shade leaves. In spite of lower ETR, higher photochemical quenching (PQ), and effective quantum yield of photosystem 2 (Φ_PS2) was evident in shade plants compared to plants exposed to higher irradiance. ETR increased linearly with increase in irradiance more under low irradiance (r ² = 0.84) compared to higher irradiance (r ² = 0.62). The maximum quantum yield of PS 2 (F_v/F_m) did not differ much in sun and shade leaves with the exception of midday when excess of light energy absorbed by plants under sun was thermally dissipated. Hence swamp taro plants adopted different strategies to utilize radiation under different irradiances. At higher irradiance, there was faster decline in proportion of open PS 2 centers (PQ) and excess light energy was dissipated through non-photochemical quenching (NPQ). Under shade, absorbed energy was effectively utilized resulting in higher Φ_PS2.     

Species-specific acclimation to strong shade modifies susceptibility of conifers to photoinhibition

Photoinhibitory processes in the photosynthetic apparatus of the seedlings of Abies alba (Mill.), Picea abies (Karst.), and Pinus mugo (Turra) growing under strong shade (5 % of full solar irradiance) or full irradiance conditions were investigated in winter and spring using chlorophyll a fluorescence techniques. The extent of photoinhibition in needles as indicated by a decrease in maximum quantum yield of PS II photochemistry (F_v/F_m) depended on species, air temperature and acclimation to the light environment. Unexpectedly, shade-tolerant Abies alba was less affected by low-temperature photoinhibition compared to the other species. F_v/F_m recovered with increasing air temperature. During winter, the seedlings of Picea abies growing in shade showed higher F_v/F_m than those from full light. Non-photochemical quenching of fluorescence (NPQ) measured at the same levels of actinic light was higher in needles acclimated to full light except for Abies alba in February. Photosynthetic performance in term of ETR (apparent electron transfer rate) was also higher in full light-acclimated needles. In April, at ambient temperature, recovery of PS II efficiency from the stress induced by illumination with saturating light was faster in the needles of Picea abies than in those of Abies alba. The shade-acclimated needles of Abies alba and Picea abies showed greater down-regulation of PS II induced by high light stress.