Leaf anatomical structures of <Emphasis Type="Italic">Paphiopedilum</Emphasis> and <Emphasis Type="Italic">Cypripedium</Emphasis> and their adaptive significance

Paphiopedilum and Cypripedium are closely related in phylogeny, but have contrasting leaf traits and habitats. To understand the divergence in leaf traits of Paphiopedilum and Cypripedium and their adaptive significance, we analyzed the leaf anatomical structures, leaf dry mass per area (LMA), leaf lifespan (LL), leaf nitrogen concentration (N _mass), leaf phosphorus concentration (P _mass), mass-based light-saturated photosynthetic rate (A _mass), water use efficiency (WUE), photosynthetic nitrogen use efficiency (PNUE) and leaf construction cost (CC) for six species. Compared with Cypripedium, Paphiopedilum was characterized by drought tolerance derived from its leaf anatomical structures, including fleshy leaves, thick surface cuticles, huge adaxial epidermis cells, lower total stoma area, and sunken stomata. The special leaf structures of Paphiopedilum were accompanied by longer LL; higher LMA, WUE, and CC; and lower N _mass, P _mass, A _mass, and PNUE compared with Cypripedium. Leaf traits in Paphiopedilum helped it adapt to arid and nutrient-poor karst habitats. However, the leaf traits of Cypripedium reflect adaptations to an environment characterized by rich soil, abundant soil water, and significant seasonal fluctuations in temperature and precipitation. The present results contribute to our understanding of the divergent adaptation of leaf traits in slipper orchids, which is beneficial for the conservation of endangered orchids.     

Litter legacy increases the competitive advantage of invasive <Emphasis Type="Italic">Phragmites australis</Emphasis> in New England wetlands

Exotic plant invaders that form monocultures and exclude native plants are often the most detrimental to native diversity and the hardest to eradicate. To generate a monoculture, the invader must garner more resources than resident natives and, once established, persist despite high densities of conspecific neighbors. Coincident with expansion and long-term persistence, successful invaders typically accumulate senesced material, but the role of this litter in mediating the invader’s ability to establish and maintain monospecific dominance has rarely been investigated. We used stands of the common reed, Phragmites australis, a prolific wetland invader in North America, to explore the impact of litter on interspecific competition with the native rush, Juncus gerardii, and intraspecific competition among live shoots. In 10 × 10 m areas positioned on Phragmites expansion fronts, we removed litter to isolate its effect from live Phragmites on light availability, aboveground biomass and community composition. Compared to adjacent, unmanipulated fronts, light availability nearly tripled and Juncus biomass increased >170% in litter removal areas after 4 months. Although the positive response of Juncus and native forbs was most pronounced on the leading edge of Phragmites stands, litter removal triggered a 271% increase in native plant biomass even in the interior of stands where Phragmites’ live stem density was highest. Litter treatment did not significantly affect Phragmites biomass, but more, shorter stems emerged in litter removals revealing Phragmites modifies stem phenotype in response to local litter and light conditions. These results suggest that litter plays a central role in Phragmites’ invasion process, from initial establishment to subsequent monospecific dominance. Thus, prescribed litter removal may be an effective strategy to enhance coexistence of native plant populations in wetlands where eradication of invasive monocultures is not an ecologically or economically feasible option.     

Biogeography of <Emphasis Type="Italic">Phragmites</Emphasis><Emphasis Type="Italic">australis</Emphasis> lineages in the southwestern United States

The environmental and social impacts of Phragmites australis invasion have been extensively studied in the eastern United States. In the West where the invasion is relatively recent, a lack of information on distributions and spread has limited our ability to manage invasive populations or assess whether native populations will experience a decline similar to that in the East. Between 2006 and 2015, we evaluated the genetic status, distribution, and soil properties (pH, electrical conductivity, and soil texture) of Phragmites stands in wetlands and riparian systems throughout the Southwest. Native (subspecies americanus), Introduced (haplotype M), and Gulf Coast (subspecies berlandieri) Phragmites lineages were identified in the survey region, as well as watershed-scale hybridization between the Native and Introduced lineages in southern Nevada. Two Asian haplotypes (P and Q) that were previously not known to occur in North America were found in California. The Native lineage was the most frequent and widespread across the region, with four cpDNA haplotypes (A, B, H, and AR) occurring at low densities in all wetland types. Most Introduced Phragmites stands were in or near major urban centers and associated with anthropogenic disturbance in wetlands and rivers, and we document their spread in the region, which is likely facilitated by transportation and urban development. Soil pH of Native and hybrid stands was higher (averaging 8.3 and 8.6, respectively) than Introduced stands (pH of 7.5) and was the only soil property that differed among lineages. Continued monitoring of all Phragmites lineages in the Southwest will aid in assessing the conservation status of Native populations and developing management priorities for non-native stands.     

Expansion of <Emphasis Type="Italic">Phragmites australis</Emphasis> (Cav.) Trin. ex Steud. (Common Reed) into <Emphasis Type="Italic">Typha</Emphasis> spp. (Cattail) Wetlands in Northwestern Indiana,USA

Expansion of Phragmites australis (Cav.) Trin. ex Steud. (common reed) into stands of Typha spp. (cattail; Typha australis L. and T. x glauca) is common in the wetlands of northwestern Indiana (USA). To understand this phenomenon better, we investigated the production of shoot sprouts and proportional allocation of biomass as well as a potential role for the water table in the relative dominance of each species. The reduction in sprouts from rhizomes upon vegetative expansion of Phragmites appeared to be the most likely process causing the decline of Typha. The latter had a shoot density of 39/m² in plots without Phragmites, but this dropped to 13 shoots m⁻² in plots that had been invaded by Phramites. Such a decline was likely caused by reduced reserves; e.g., the belowground biomass of Typha decreased from 11.3 g m⁻² without Phragmites to 8.1 g m⁻² with Phragmites. The latter also reduced its belowground biomass but not its shoot density in the presence of Typha. The mean weight of Phragmites shoots was 2.9 g, and nearly all produced inflorescences. Meanwhile, Typha failed to develop spadices despite its shoots having a greater biomass (7 g). This suggests that Phragmites is more efficient than Typha in shoot growth. Springtime flooding appeared to promote the sprout of Typha shoots from shallow rhizomes (≈18 cm below the soil surface), whereas the shoot density of Phragmites showed no correlation with water level in that season. Deep-rooted Phragmites (≈39 cm) occurred on both high and low water-table sites, whereas the shallow-rooted Typha was limited to only the former. Phragmites will likely continue its expansion, by vegetative sprouts from rhizomes, into Typha wetlands.     

Photosynthetic performance of the aquatic macrophyte <Emphasis Type="Italic">Althenia orientalis</Emphasis> to solar radiation along its vertical stems

We have studied the plasticity of the photosynthetic apparatus in the endangered aquatic macrophyte Althenia orientalis to the gradient of light availability within its meadow canopy. We determined diurnal change in situ irradiance, light quality, in vivo chlorophyll a fluorescence, ex situ oxygen evolution rates, respiration rate and pigment concentration. The levels of photosynthetic photon flux density (PFD) and ultraviolet radiation (UVR) and the red/far-red ratio decreased with depth within the canopies of A. orientalis. Apical leaves had a greater decrease of the maximal quantum yield (F _v/F _m) in the morning and a faster recovery rate in the afternoon than those in the basal ones. The relative electron transport rate (ETRr) was not saturated at any time of the day, even in the apical leaves that received the highest light. The maximum light-saturated rate of gross photosynthesis (GP_max) took place in apical leaves around noon. The chlorophyll a/b ratio values were higher, and the chlorophyll/carotenoid ratio values lower, in apical leaves than basal ones. The highest concentrations in total carotenoids were reached in the apical leaves around noon. A. orientalis has a high capacity to acclimatize to the changes in the light environment, both in quality and quantity, presenting sun and shade leaves in the same stem through the vertical gradient in the canopy.     

Leaf traits of natural populations of <Emphasis Type="Italic">Adiantum reniforme</Emphasis> var. <Emphasis Type="Italic">sinensis</Emphasis>, endemic to the Three Gorges region in China

Leaf mass per unit area (LMA), carbon and nitrogen contents, leaf construction cost, and photosynthetic capacity (P _max) of Adiantum reniforme var. sinensis, an endangered fern endemic to the Three Gorges region in southwest China, were compared in five populations differing in habitat such as soil moisture and irradiance. The low soil moisture and high irradiance habitat population exhibited significantly higher LMA, area-based leaf construction (CC_A), and carbon content (C_A), but lower leaf nitrogen content per unit dry mass (N_M) than the other habitat populations. The high soil moisture and low irradiance habitat populations had the lowest CC_A, but their cost/benefic ratios of CCA/P _max were similar to the medium soil moisture and irradiance habitat population due to their lower leaf P _max. Hence A. reniforme var. sinensis prefers partially shaded, moist but well-drained, slope habitats. Due to human activities, however, its main habitats now are cliffs or steeply sloped bare rocks with poor and thin soil. The relatively high energy requirements and low photosynthetic capacity in these habitats could limit the capability of the species in extending population or interspecific competition and hence increase its endangerment.     

Spatial and temporal variations in the photosynthesis-nitrogen relationship in a Japanese cedar (<Emphasis Type="Italic">Cryptomeria japonica</Emphasis> D. Don) canopy

Spatial and temporal variations in light-saturated photosynthetic capacity and needle nitrogen (N) content were investigated in one 8 m tall Japanese cedar (Cryptomeria japonica D. Don) canopy for a full year. The photosynthetic capacity and needle N content in various layers of the canopy were measured every month. Temporal variations in photosynthetic capacity and needle N content expressed on a projected-area basis (P_area, N_area) were similar. Furthermore, both P_area and N_area decreased with increasing depth from the top of the canopy on each sampling date. As a consequence, a significant correlation was observed between N_area and P_area. Temporal variations in photosynthetic capacity and needle N content expressed on a mass basis (P_mass, N_mass) were also similar. P_mass also decreased with increasing canopy depth. However, in contrast to N_area, there was only a slight decrease in N_mass with increasing canopy depth. Hence, the correlation between N_mass and P_mass was lower than the projected-area value. Because N_area was highly correlated with the needle mass per projected-area (NMA), the spatial variation in N_area (and therefore P_area) in the canopy is attributed to the variation in NMA, which decreases as the depth from the top of the canopy increases. Furthermore, the slope of the linear regression between N_area and NMA differed between sampling dates, indicating that the temporal variations in N_area (and therefore P_area) are strongly influenced by N_mass. For most of the sampling dates, a linear regression between N_area and P_area tends to converge into a single line segment. However, on several sampling dates, there was a pronounced decline in P_area below this line segment. This reduction in P_area, which does not accompany a reduction in Narea, seems to be attributable to stomatal limitations induced by the low soil temperature in winter and early spring.     

Improvement of growth conditions and gas exchange of<Emphasis Type="Italic"> Fagus sylvatica</Emphasis> L. seedlings planted below a recently thinned<Emphasis Type="Italic"> Pinus sylvestris</Emphasis> L. stand

Abstract Gas exchange and growth of beech seedlings planted in the understory of a recently thinned pinewood were recorded for 2 years. Relative irradiance was assessed by hemispherical photographs taken just after the thinning. Predawn water potential (_pd), daily gas exchange and chlorophyll fluorescence were measured several times during the two growing seasons. Maximum values of photosynthesis (A _max) and stomatal conductance to water vapour (g _wvmax) were established from daily data. Maximum quantum efficiency of PS II was recorded at dawn by taking the variable to maximum chlorophyll fluorescence ratio on dark adapted leaves (F _v/F _m). In the middle of each summer, leaf nitrogen content and leaf mass per area were evaluated, and height growth and basal area increment were recorded at the end of the season. The thinning treatment removed half the trees and generated around 10% more available relative irradiance (GLF). This was followed by an increase in net photosynthesis at saturating PPFD (A _sat) and in maximum stomatal conductance to water vapour (g _wvmax). Moreover, specific leaf mass (SLM) and mass based nitrogen content (N_m) showed higher values for seedlings in the thinned stand. In both years, a positive relationship was established between the area based nitrogen content (N_a) and maximum net photosynthesis (A _max). In 1998, a year with a dry summer, seedlings suffered a significant drop in daily A _max irrespective of the thinning regime. This was a response to an increase in stomatal limitation to net photosynthesis, g _wvmax reaching the lowest value on dates with the highest drought. A lack of decrease of Fv/Fm confirmed the absence of significant non-stomatal limitation to A as a consequence of photoinhibition after opening the pinewood. A higher maximum quantum efficiency of open PS II centres (Fv/Fm) was registered in seedlings in the thinned stand. The significance of the differences between the treatments was stronger in the second year after thinning. In 1999, a year with frequent summer storms, water availability increased for seedlings growing under the thinned pinewood. Overall, the reduced pine overstory had a positive effect on physiological responses of beech seedlings, which was translated into improved seedling growth.     

Complex invader-ecosystem interactions and seasonality mediate the impact of non-native <Emphasis Type="Italic">Phragmites</Emphasis> on CH<Subscript>4</Subscript> emissions

Invasive plants can influence ecosystem processes such as greenhouse gas (GHG) emissions from wetland systems directly through plant-mediated transfer of GHGs to the atmosphere or through indirect modification of the environment. However, patterns of plant invasion often co-vary with other environmental gradients, so attributing ecosystem effects to invasion can be difficult in observational studies. Here, we assessed the impact of Phragmites australis invasion into native shortgrass communities on methane (CH₄) emissions by conducting field measurements of CH₄ emissions along transects of invasion by Phragmites in two neighboring brackish marsh sites and compared these findings to those from a field-based mesocosm experiment. We found remarkable differences in CH₄ emissions and the influence of Phragmites on CH₄ emissions between the two neighboring marsh sites. While Phragmites consistently increased CH₄ emissions dramatically by 10.4 ± 3.7 µmol m^?2 min^?1 (mean ± SE) in our high-porewater CH₄ site, increases in CH₄ emissions were much smaller (1.4 ± 0.5 µmol m^?2 min^?1) and rarely significant in our low-porewater CH₄ site. While CH₄ emissions in Phragmites-invaded zones of both marsh sites increased significantly, the presence of Phragmites did not alter emissions in a complementary mesocosm experiment. Seasonality and changes in temperature and light availability caused contrasting responses of CH₄ emissions from Phragmites- versus native zones. Our data suggest that Phragmites-mediated CH₄ emissions are particularly profound in soils with innately high rates of CH₄ production. We demonstrate that the effects of invasive species on ecosystem processes such as GHG emissions may be predictable qualitatively but highly variable quantitatively. Therefore, generalizations cannot be made with respect to invader-ecosystem processes, as interactions between the invader and local abiotic conditions that vary both spatially and temporally on the order of meters and hours, respectively, can have a stronger impact on GHG emissions than the invader itself.     

Dynamics of leaf area and nitrogen in the canopy of an annual herb, <Emphasis Type="Italic">Xanthium canadense</Emphasis>

We studied leaf area and nitrogen dynamics in the canopy of stands of an annual herb Xanthium canadense, grown at a high (HN)- and a low-nitorgen (LN) availability. Standing leaf area increased continuously through the vegetative growth period in the LN stand, or leveled off in the later stage in the HN stand. When scaled against standing leaf area, both production and loss rates of leaf area increased but with different patterns: the production rate was retarded, while the loss rate was accelerated, implying an upper limit of standing leaf area of the canopy. The rate of leaf-area production was higher in the HN than in the LN stand, which was caused by the higher rate of leaf production per standing leaf area as well as the greater standing leaf area in the HN stand. Although the rate of leaf-area loss was higher in the HN than in the LN stand, it was not significantly different between the two stands when compared at a common standing leaf area, suggesting involvement of light climate in determination of the leaf-loss rate. On the other hand, the rate of leaf-area loss was positively correlated with nitrogen demand for leaf area development across the two stands, suggesting that leaf loss was caused by retranslocation of nitrogen for construction of new leaves. A simple simulation model of leaf and nitrogen dynamics in the canopy showed that, at steady state, where the rate of leaf-area loss becomes equal to the production rate, the standing leaf area was still greater in the HN than in the LN stand. Similarly, when the uptake and loss of nitrogen are equilibrated, the standing nitrogen was greater in the HN than in the LN stand. These results suggest that leaf-area production is strongly controlled by nitrogen availability, while both nitrogen and light climate determine leaf-loss rates in the canopy.     

Understanding invasion as a process: the case of <Emphasis Type="Italic">Phalaris arundinacea</Emphasis> in wet prairies

Invasive plants that most threaten biodiversity are those that rapidly form a monospecific stand, like the clonal grass, Phalaris arundinacea. Understanding complex and potentially interacting factors that are common in urban and agricultural landscapes and underlie rapid invasions requires an experimental, factorial approach. We tested the effects of flooding and nutrient and sediment additions (3 × 3 × 3 = 27 treatments, plus a control with no additions) on invasion of Phalaris into mesocosms containing wet prairie vegetation. We discovered a three-step invasion and degradation process: (1) initially, resident native species declined with prolonged flooding and sediment additions, and (2) prolonged flooding, sedimentation, and nutrients accelerated Phalaris aboveground growth; biomass rose to 430 times that of the control within just two growing seasons. The dramatic expansion of Phalaris in the second year resulted in the formation of monospecific stands in over one-third of the treatments, as (3) native species continued their decline in year 2. Disturbances acted alone and in combination to make the resident wetland community more invasible and Phalaris more aggressive, leading to monospecific stands. Yet, Phalaris did not always “win”: under the least disturbed conditions, the resident plant canopy remained dense and vigorous and Phalaris remained small. When anthropogenic disturbances coincide with increases in the gross supply of resources, more tolerant, fast-growing, and morphologically plastic plants like Phalaris can invade very rapidly. The fluctuating resource hypothesis should thus be refined to consider the role of interacting disturbances in facilitating invasions.     

PhragNet: crowdsourcing to investigate ecology and management of invasive <Emphasis Type="Italic">Phragmites australis</Emphasis> (common reed) in North America

Invasion biology research, often performed by scientists at relatively small spatial scales, provides experimental precision but may be limited in generalizability. Conversely, large-scale invasive species management represents a largely untapped wealth of information on invasion ecology and management, but such data are difficult to capture and synthesize. We developed a network (“PhragNet”) of individuals managing wetlands occupied by native and non-native lineages of the invasive wetland grass Phragmites australis (common reed). This network collected environmental and genetic samples, habitat data, and management information to identify environmental and plant community associations of Phragmites invasion and patterns of management responses. Fifty managers overseeing 209 Phragmites stands in 16 US states and ON, Canada participated. Participants represented federal agencies (26%), municipalities (20%), NGOs (20%), academia (14%), state agencies (12%), and private landowners (8%). Relative to the native lineage, non-native Phragmites occurred in areas with higher nitrate/nitrite and ammonium than non-native Phragmites. Stand interiors had higher soil electrical conductivity than nearby uninvaded areas, consistent with use of road salt promoting spread of Phragmites. Non-native Phragmites co-occurred with fewer plant species than native Phragmites and was actively targeted for management. Herbicide was applied to 51% of non-native stands; surprisingly, 11% of native stands were also treated with herbicide. This project demonstrates the utility of crowdsourcing standardized data from resource managers. We conclude by describing how this approach could be expanded into an adaptive management framework, strengthening connections between wetland management and research.     

Functional correlates of leaf demographic response to gap release in saplings of a shade-tolerant tree,<Emphasis Type="Italic"> Elateriospermum tapos</Emphasis>

For a shade-tolerant SE Asian tropical tree, Elateriospermum tapos (Euphorbiaceae), we studied field-established saplings in gaps and the shaded understory to test the hypothesis that differences in leaf demography and leaf life span under contrasting light regimes should be functionally correlated with architecture, self-shading and nitrogen distribution within the sapling crown. Rates of leaf production and net leaf gain were greater for saplings in gaps than those in the understory. Median leaf life span was approximately 26 months in the gap saplings, while it was estimated to be greater than 38 months in the understory saplings. Consequently, gap saplings had a greater standing leaf number and experienced greater degrees of self-shading than understory saplings. Light availability at individual leaves, estimated by a combination of canopy photos and a three-dimensional architecture model, were negatively correlated with leaf age in gap saplings but not so in understory saplings. Leaf nitrogen content per unit area (N_area) was influenced more by light availability than by leaf age in the gap saplings. In contrast, in understory saplings, N_area was neither correlated with light availability nor with leaf age, and did not decrease significantly before 38 months in leaf age. We conclude that saplings of this shade-tolerant species apparently prolong their leaf life span in the shaded understory through slower rates of leaf production, lower standing number of leaves and lower degrees of self shading than in gap, and that the rate of decline of N_area with leaf age depends on architecture and self-shading regimes that respond to changes in light regimes.     

The physiological advantage of an ecological filter species, <Emphasis Type="Italic">Indocalamus longiauritus</Emphasis>, over co-occurring <Emphasis Type="Italic">Fagus lucida</Emphasis> and <Emphasis Type="Italic">Castanopsis lamontii</Emphasis> seedlings

Indocalamus longiauritus (a dwarf bamboo) dominates forest understory and functions as an ecological filter to hinder the regeneration of canopy tree species in many temperate forests. However, the physiological mechanism underlying the function of ecological filters is not clear. In this study, we measured leaf-level carbon capture ability and use efficiency of the dwarf bamboo and the co-existing Fagus lucida (beech) and Castanopsis lamontii (chinkapin) seedlings in forest understory and small gaps in a beech–chinkapin mixed forest in the summer of 2005. The results indicated that I. longiauritus exhibited greater carbon capture ability, as indexed by light-saturated photosynthetic rate (P _max), maximal carboxylation rate, maximal electron transport rate and carboxylation efficiency, than the co-occurring F. lucida and C. lamontii seedlings in both forest understory and small gaps. Higher carbon capture ability in I. longiauritus was related to its greater partition of absorbed light energy to photochemistry. I. longiauritus had higher photosynthetic nitrogen use efficiency than F. lucida and C. lamontii seedlings in both light environments. However, water use efficiency (WUE) in I. longiauritus was higher than F. lucida but lower than C. lamontii. This intermediate WUE in I. longiauritus was related to its intermediate light-saturated stomatal conductance. In addition, I. longiauritus reduced metabolic cost by increasing the ratio of P _max to respiration rate, leading to increased net carbon balance. On the other hand, F. lucida and C. lamontii seedlings had greater plasticity of carbon capture ability and leaf structural traits, which might facilitate colonization of gaps and realization of natural regeneration in these species.     

Effects of <Emphasis Type="Italic">Morella faya</Emphasis> tree invasion on aboveground carbon storage in Hawaii

The negative effects of biological invasion are often the focus of ecological studies, but few have considered potential positive impacts, such as increased carbon storage, resulting from invasion. We combined airborne imaging spectrometer and LiDAR (light detection and ranging) observations with field measurements to assess if the highly invasive nitrogen-fixing tree Morella faya alters canopy 3-D structure and aboveground biomass (AGB) along a 1,500 mm precipitation gradient in Hawaii. Airborne analysis of canopy water content, leaf nitrogen concentration, fractional canopy cover, and vegetation height facilitated mapping of native- and Morella-dominated canopies in rainforest, woodland–savanna and shrubland ecosystems, with Morella detection errors ranging from 0 to 13.4%. Allometric equations were developed to relate the combined LiDAR and spectral data to field-based AGB estimates (r ² = 0.97, P < 0.01), and to produce a map of biomass stocks throughout native and invaded ecosystems. The structure of the invasive Morella canopies varied by ecosystem type, and the invader shaded out native understory plants in rainforest zones. Despite a 350% increase in AGB going from shrubland to rainforest, Morella did not increase average AGB in any ecosystem it invaded. Furthermore, spatial distributions of biomass indicated that Morella decreased maximum AGB in the woodland–savanna ecosystems. We conclude that Morella tree invasion does not enhance aboveground carbon stocks in any ecosystem it invades in Hawaii, thereby minimizing its contribution to this potentially important ecosystem service. We also found that the fusion of spectral and LiDAR remote sensing provided canopy chemical and structural data facilitating a landscape assessment of how biological invasion alters on carbon stocks and other ecosystem properties.     

Photosynthetic responses to short-term and long-term light variation in <Emphasis Type="Italic">Pinus sylvestris</Emphasis> and <Emphasis Type="Italic">Salix dasyclados</Emphasis>

Pinus sylvestris and Salix dasyclados, which differ in leaf longevity, were compared with respect to four aspects of photosynthetic light use and response: high light acclimation, photoinhibition resistance and recovery, lightfleck exposure and use and chloroplast acclimation across leaves. The first two aspects were examined using seedlings under controlled conditions and the other two were tested using trees in the field. When exposed to high light, shade leaves of Pinus acclimated completely, achieving the same photosynthetic capacities as sun leaves, whereas shade leaves of Salix did not reach sun leaf capacities although the absolute magnitude of their acclimation was larger. Shade leaves of Pinus were also more resistant to photoinhibition than those of Salix. Much of the direct light supplied within the canopy was in the form of rapid fluctuations, lightflecks, for Pinus and Salix alike. They exploited short lightflecks with similar efficiency. The greater proportion of diffuse light in the canopy for Pinus than Salix seems to lead to a lesser degree of differential intra-leaf acclimation of chloroplasts, in turn leading to lower efficiency of photosynthesis under unilateral light as reflected by a lower convexity, rate of bending, of the light–response curve. The differences in light use and responses are discussed in relation to possible differences in characteristics of the long and short-lived leaf.     

Leaf-trait relationships of <Emphasis Type="Italic">Quercus liaotungensis</Emphasis> along an altitudinal gradient in Dongling Mountain,Beijing

Few studies have examined leaf-trait relationships in the distribution of individual species along an environmental gradient. Here we address the issue by testing for the leaf-trait relationships of Quercus liaotungensis, a dominant deciduous woody species in northern China, along an altitudinal gradient in Dongling Mountain, Beijing. These leaf traits included specific leaf area (SLA), leaf dry matter content (LDMC), and leaf nitrogen, phosphorus, and potassium concentration on mass basis (N_mass, P_mass and K_mass, respectively). Along the altitudinal gradient, negative relationships between SLA and LDMC and N_mass were found, and N_mass, P_mass and K_mass correlated with each other positively. Relationship between N_mass and P_mass was stronger than the ones between N_mass and K_mass, and between P_mass and K_mass. The weak and negative relationship between SLA and N_mass might result from trade-offs that limit photosynthesis and water use efficiency along the altitudinal gradient, suggesting many environmental factors of local site being the collective forcing of drivers. Out of our expectations, N_mass and P_mass related very weakly to soil N and P, and no significant relationship between K_mass and soil K was found along elevation. These relationships could be used to predict the productivity of the population with changing environment in this region.     

Tissue culture and wetland establishment of the freshwater monocots <Emphasis Type="Italic">Carex,Juncus, Scirpus</Emphasis>, and <Emphasis Type="Italic">Typha</Emphasis>

Summary Cell cultures of freshwater wetland monocots were regenerated, plants were grown in the greenhouse, and then established and evaluated in wetlands. Typha (cattail), Juncus (rushes), Scirpus (bulrushes), and Carex (sedges) were studied because they are common, dominant, high biomass wetland-adapted plants, tolerant of chemically diverse ecosystems. The goal was to define micropropagation and wetland establishment protocols. Tissue culture systems defined for numerous monocot crop species can be readily applied to wetland plants, with a few modifications. Issues addressed were selection of explant material, shoot and root regeneration conditions, culture age verses regenerability, greenhouse acclimatization needs, plant uniformity and requirements for wetland establishment. In vitro-germinated seedlings were an excellent source of pathogen-free regenerable tissue. T. latifolia, T. angustifolia, and J. accuminatus were regenerated from callus induced in the dark with picloram, then transferred to medium with benzyladenine in the light to promote shoot organogenesis. J. effusus, S. polyphyllus, and C. lurida could not be regenerated from callus, which turned black. They could be regenerated directly by culturing intact seedlings directly on cytokinin media in the light. Shoots rooted with little or no auxin. J. effusus rooting was promoted by the addition of charcoal to the medium. Covering plants for the first 2 wk with plastic facilitated greenhouse establishment. There were high rates of greenhouse and wetland survival. No abnormal plants were observed. These regeneration systems could be utilized for the production of wetland plants for potential application in habitat restoration and wetland creation, and would provide an alternative to field collection.     

Characteristics of leaf photosynthesis and simulated individual carbon budget in <Emphasis Type="Italic">Primula nutans</Emphasis> under contrasting light and temperature conditions

Primula nutans Georgi is widely distributed in hummock-and-hollow wetlands on the Qinghai-Tibetan Plateau. To assess the ecophysiology of this species in responding to microenvironments, we examined the photosynthetic characteristics and individual carbon gain of plants growing in different microsites from a hummock-and-hollow wetland on the Qinghai-Tibetan Plateau and under laboratory conditions. Plants from wetland hummock microsites showed significantly higher light-saturated photosynthetic CO₂ uptake (A _max) than those from microsites in hollows at a controlled temperature of 15°C in leaf chamber. Leaf dark respiration rate (R) was only significantly higher in plants from hummocks than hollows at the measuring temperature of 35°C. Optimum temperature for A _max was 15°C for all plants in the field despite different microsites. In plants growing under laboratory conditions differing in light and temperature, both A _max and R were significantly higher under higher growth light (photosynthetic photon flux density, PPFD: 800 or 400 μmol m⁻² s⁻¹) than low light of 90 μmol m⁻² s⁻¹. No statistically significant differences in A _max and R existed in plants differing in growing temperatures. Estimates derived from the photosynthetic parameters of field plants, and microsite environmental measures including PPFD, air temperature and soil temperature showed that the optimum mean daily temperature for net daily carbon gain was around 10°C and the net daily carbon gain was largely limited under lower daily total PPFD. These results suggest that the differences in A _max and R in P. nutans are strongly affected by growing light regimes but not by temperature regimes. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Photosynthetic performance of <Emphasis Type="Italic">Lycoris radiata</Emphasis> var. <Emphasis Type="Italic">radiata</Emphasis> to shade treatments

The effects of shade on the growth, leaf photosynthetic characteristics, and chlorophyll (Chl) fluorescence parameters of Lycoris radiata var. radiata were determined under differing irradiances (15, 65, and 100% of full irradiance) within pots. The HI plants exhibited a typical decline in net photosynthetic rate (P _N) during midday, which was not observed in MI- and LI plants. This indicated a possible photoinhibition in HI plants as the ratio of variable to maximum fluorescence (F_v/F_m) value was higher and the minimal fluorescence (F₀) was lower in the, and LI plants. Diurnal patterns of stomatal conductance (g _s) and transpiration rate (E) were remarkably similar to those of P _N at each shade treatments, and the intercellular CO₂ concentration (C _i) had the opposite change trend. Under both shading conditions, the light saturation point, light compensation point and photon-saturated photosynthetic rate (P _max) became lower than those under full sunlight, and it was the opposite for the apparent quantum yield (AQY). The higher the level of shade, the lower the integrated daytime carbon gain, stomatal and epidermis cell densities, specific leaf mass (SLM), bulb mass ratio (BMR), leaf thickness, and Chl a/b ratio. In contrast, contents of Chls per dry mass (DM), leaf area ratio (LAR), leaf mass ratio (LMR), leaf length, leaf area and total leaf area per plant increased under the same shade levels to promote photon absorption and to compensate for the lower radiant energy. Therefore, when the integrated daytime carbon gain, leaf area and total leaf area per plant, which are the main factors determining the productivity of L. radiata var. radiata plant, were taken into account together, this species may be cultivated at about 60∼70% of ambient irradiance to promote its growth.