Effect of Hydrologic Restoration and Lonicera maackii Removal on Herbaceous Understory Vegetation in a Bottomland Hardwood Forest

Amur honeysuckle (Lonicera maackii (Rupr.) Herder), a large deciduous shrub from China, has invaded many forests in eastern/central United States. The species was removed by cutting and herbicide application from a recently hydrologically restored section of a bottomland hardwood forest in central Ohio, and the response of understory plants, especially herbaceous species, was measured. Plots were established in uncleared and cleared sections, and percent cover of each herbaceous understory species was estimated monthly. One season after several years of Lonicera removal efforts, no significant association was discovered between percentage of Lonicera cover and total understory species abundance. There was, however, a direct correlation between elevation and honeysuckle abundance; L. maackii abundance was negatively associated with low elevations, likely due to hydrologic factors. Plant species diversity (H) and richness (s) increased with elevation but were not significantly different on plots with honeysuckle removal (H = 0.86 ± 0.08 vs. 0.78 ± 0.09 and s = 4.4 ± 0.19 vs. 4.2 ± 0.2 species/m², respectively) despite the fact that understory light levels measured by densiometer were significantly higher (α= 0.003) in cleared versus uncleared sections. Native and invasive species were found in similar proportions in the two sections, and significant sprouting and regrowth of L. maackii were observed throughout the cleared section. Although the removal of L. maackii altered the characteristics of the plant species assemblage, the value of this management remains questionable in the years immediately following treatment.     

EFFECTS OF CO2 ENRICHMENT ON THE GROWTH AND MORPHOLOGY OF A NATIVE AND AN INTRODUCED HONEYSUCKLE VINE

Japanese honeysuckle (Lonicera japonica Thunb.), introduced to the United States, and the native coral honeysuckle (Lonicera sempervirens L.) were compared to determine how intrinsic differences in their growth characteristics would affect their response to atmospheric carbon dioxide enrichment. Plants of both species grown from cuttings were harvested after 54 days of growth in controlled environment growth chambers at 350, 675, or 1,000 μl/liter CO₂. The biomass of Japanese honeysuckle was increased 135% at 675 μ∗∗∗l/liter CO₂ and 76% at 1,000 μl/liter CO₂ after 54 days. Morphologically, the main effect of CO₂ enrichment was to triple the number of branches and to increase total branch length six times. Enhanced and accelerated branchingalso increased total leaf area 50% at elevated CO₂ concentrations. In coral honeysuckle, total biomass was only 40% greater in the elevated CO₂ treatments. Branching was quadrupled but had not proceeded long enough to affect total leaf area. Main stem height was increased 36% at 1,000 μl/liter CO₂. The much less significant height response of other woody erect growth forms suggests that vines may increase in importance during competition if atmospheric CO₂ concentrations increase as predicted. The impact of Japanese honeysuckle in the United States may become more serious.     

Links between Belowground and Aboveground Resource-Related Traits Reveal Species Growth Strategies that Promote Invasive Advantages

Belowground processes are rarely considered in comparison studies of native verses invasive species. We examined relationships between belowground fine root production and lifespan, leaf phenology, and seasonal nitrogen dynamics of Lonicera japonica (non-native) versus L. sempervirens (native) and Frangula alnus (non-native) versus Rhamnus alnifolia (native), over time. First and second order fine roots were monitored from 2010 to 2012 using minirhizotron technology and rhizotron windows. ¹⁵N uptake of fine roots was measured across spring and fall seasons. Significant differences in fine root production across seasons were seen between Lonicera species, but not between Frangula and Rhamnus, with both groups having notable asynchrony in regards to the timing of leaf production. Root order and the number of root neighbors at the time of root death were the strongest predictors of root lifespan of both species pairs. Seasonal ¹⁵N uptake was higher in spring than in the fall, which did not support the need for higher root activity to correspond with extended leaf phenology. We found higher spring ¹⁵N uptake in non-native L. japonica compared to native L. sempervirens, although there was no difference in ¹⁵N uptake between Frangula and Rhamnus species. Our findings indicate the potential for fast-growing non-native Lonicera japonica and Frangula alnus to outcompete native counterparts through differences in biomass allocation, root turnover, and nitrogen uptake, however evidence that this is a general strategy of invader dominance is limited.     

Interactions between two co-dominant, invasive plants in the understory of a temperate deciduous forest

Negative interactions between non-indigenous and native species has been an important research topic of invasion biology. However, interactions between two or more invasive species may be as important in understanding biological invasions, but they have rarely been studied. In this paper, we describe three field experiments that investigated interactions between two non-indigenous plant species invasive in the eastern United States, Lonicera japonica (a perennial vine) and Microstegium vimineum (an annual grass). A press removal experiment conducted within a deciduous forest understory community indicated that M. vimineum was a superior competitor to L. japonica. We tested the hypothesis that the competitive success of M. vimineum was because it overgrew, and reduced light available to, L. japonica, by conducting a separate light gradient experiment within the same community. Shade cloth that simulated the M. vimineum canopy reduced the performance of L. japonica. In a third complementary experiment, we added experimental support hosts to test the hypothesis that the competitive ability of L. japonica is limited by support hosts, onto which L. japonica climbs to access light. We found that the abundance of climbing branches increased with the number of support hosts. Results of this experiment indicate that these two invasive species compete asymmetrically for resources, particularly light.     

Effects of Irradiance during Growth on Adaptive Photosynthetic Characteristics of Velvetleaf and Cotton

We grew velvetleaf (Abutilon theophrasti Medic.) and cotton (Gossypium hirsutum L. var. Stoneville 213) at three irradiances and determined the photosynthetic responses of single leaves to a range of six irradiances from 90 to 2000 μeinsteins m⁻²sec⁻¹. In air containing 21% O₂, velvetleaf and cotton grown at 750 μeinsteins m⁻²sec⁻¹ had maximum photosynthetic rates of 18.4 and 21.9 mg of CO₂ dm⁻²hr⁻¹, respectively. Maximum rates for leaves grown at 320 and 90 μeinsteins m⁻²sec⁻¹ were 15.3 and 10.3 mg of CO₂ dm⁻²hr⁻¹ in velvetleaf and 12 and 6.7 mg of CO₂ dm⁻²hr⁻¹ in cotton, respectively. In 1 O₂, maximum photosynthetic rates were 1.5 to 2.3 times the rates in air containing 21% O₂, and plants grown at medium and high irradiance did not differ in rate. In both species, stomatal conductance was not significantly affected by growth irradiance. The differences in maximum photosynthetic rates were associated with differences in mesophyll conductance. Mesophyll conductance increased with growth irradiance and correlated positively with mesophyll thickness or volume per unit leaf area, chlorophyll content per unit area, and photosynthetic unit density per unit area. Thus, quantitative changes in the photosynthetic apparatus help account for photosynthetic adaptation to irradiance in both species. Net assimilation rates calculated for whole plants by mathematical growth analysis were closely correlated with single-leaf photosynthetic rates.     

Exotic Lonicera species both escape and resist specialist and generalist herbivores in the introduced range in North America

The enemy release hypothesis predicts that invasive plant species may benefit from a lack of top-down control by co-evolved herbivores, particularly specialists, in their new range. However, to benefit from enemy escape, invasive plants must also escape or resist specialist or generalist herbivores that attack related species in the introduced range. We compared insect herbivore damage on the exotic shrub, Lonicera maackii, the native congener Lonicera reticulata, and the native confamilial Viburnum prunifolium in North America. We also compared the laboratory preference and performance of a North American honeysuckle specialist sawfly (Zaraea inflata) and the performance of a widespread generalist caterpillar (Spodoptera frugiperda) on cut foliage from native and exotic Lonicera species. L. maackii received significantly lower amounts of foliar herbivory than L. reticulata across three seasons, while damage levels observed on V. prunifolium for two seasons was generally intermediate between L. reticulata and L. maackii. The specialist sawfly damaged L. reticulata heavily, but was not detected on L. maackii in the field. There were few statistical differences in the performance of sawfly larvae on L. reticulata and L. maackii, but the sawfly achieved higher pupal masses on L. reticulata than on L. maackii, and they strongly preferred L. reticulata over L. maackii when given a choice. The sawfly was unable to complete development on native L. sempervirens and non-native L. japonica. In contrast, the generalist caterpillar performed similarly on all Lonicera species. While L. maackii experienced little herbivory in the field compared to native relatives in the same habitat, laboratory assays indicate L. maackii appears to be a suitable host that escapes selection by the specialist, but L. japonica and L. sempervirens are highly resistant to it. These findings indicate that both enemy escape and resistance (to a specialist, but not a generalist herbivore) may contribute to the success of exotic Lonicera species.     

Scaling carbon dioxide and water vapour exchange from leaf to canopy in a deciduous forest. I. Leaf model parametrization

In order to parametrize a leaf submodel of a canopy level gas-exchange model, a series of photosynthesis and stomatal conductance measurements were made on leaves of white oak (Quercus alba L.) and red maple (Acer rubrum L.) in a mature deciduous forest near Oak Ridge, TN. Gas-exchange characteristics of sun leaves growing at the top of a 30 m canopy and of shade leaves growing at a depth of 3–4 m from the top of the canopy were determined. Measured rates of net photosynthesis at a leaf temperature of 30°C and saturating photosynthetic photon flux density, expressed on a leaf area basis, were significantly lower (P = 0.01; n = 8) in shade leaves (7.9μmol m^?2 s^?1) than in sun leaves (11–5μmol m^?2 s^?1). Specific leaf area increased significantly with depth in the canopy, and when photosynthesis rates were expressed on a dry mass basis, they were not significantly different for shade and sun leaves. The percentage leaf nitrogen did not vary significantly with height in the canopy; thus, rates expressed on a per unit nitrogen basis were also not significantly different in shade and sun leaves. A widely used model integrating photosynthesis and stomatal conductance was parametrized independently for sun and shade leaves, enabling us to model successfully diurnal variations in photosynthesis and evapotranspiration of both classes of leaves. Key photosynthesis model parameters were found to scale with leaf nitrogen levels. The leaf model parametrizations were then incorporated into a canopy-scale gas-exchange model that is discussed and tested in a companion paper (Baldocchi & Harley 1995, Plant, Cell and Environment 18, 1157–1173).     

Duration of freezing necessary to damage the leaves of Fallopia japonica (Houtt.) Ronse Decraene

Fallopia japonica (Houtt.) Ronse Decraene is an invasive plant species that introduces economic, social, and environmental stresses. After observing frost damage to F. japonica plants in the field, we exposed leaves of F. japonica and a native species (Acer saccharum Marshall) to freezing temperatures in the laboratory and compared their net photosynthetic rate to that of fresh leaves. In both species, the net photosynthetic rate of leaves frozen for 0.5 h or for 1 h were not significantly different from each other but were both significantly less than that of fresh leaves. Fresh leaves of F. japonica had a higher net photosynthetic rate than those of A. saccharum, but the relationship was reversed in all freezing treatments. Frozen leaves of F. japonica contained microscopically visible frost lenses, which revealed the mechanism of the damage. These results quantify how quickly F. japonica is damaged by freezing conditions and suggest that minimum vernal temperatures may limit its range expansion.     

Effects of below- and aboveground competition from the vines Lonicera japonica and Parthenocissus quinquefolia on the growth of the tree host Liquidambar styraciflua

Detrimental effects of vines on tree growth in successional environments have been frequently reported. Little is known, however, about the relative importance of below and aboveground competition from vines on tree growth. The objective of this study was to quantify and compare the growth responses of Liquidambar styraciflua saplings to below and/or aboveground competition with the exotic evergreen vine, Lonicera japonica (Japanese honeysuckle), and the native deciduous vine, Parthenocissus quinquefolia (Virginia creeper). Soil trenching and/or vine-trellising were used to control the type of vine competition experienced by trees. Comparisons among untrenched treatments tested for effects of belowground competition. Comparisons among trenched treatments tested for effects of aboveground competition. After two growing seasons, Lonicera japonica had a greater effect on the growth of L. styraciflua than did P. quinquefolia. This effect was largely due to root competition, as canopy competition only had a negative effect on tree growth when it occurred in combination with root competition. Leaf expansion was consistently and similarly affected by all treatments which involved belowground competition.     

Photosynthetic and biomass allocation responses of Liquidambar styraciflua (Hamamelidaceae) to vine competition

This 2-year field study examined stomatal conductance, photosynthesis, and biomass allocation of Liquidambar styraciflua saplings in response to below- and aboveground competition with the vines Lonicera japonica and Parthenocissus quinquefolia. Vine competition did not affect stomatal conductance of the host trees. The leaf photosynthetic capacity and photosynthetic nitrogen-use efficiency were significantly reduced by root competition with vines, either singly or in combination with aboveground competition, early in the second growing season. However, such differences disappeared by the end of the second growing season. Trees competing below ground with vines also had lower allocation to leaves compared with steins. Aboveground competition with vines resulted in reduced photosynthetic capacity per unit leaf area, but not per unit leaf weight, in trees. No correlation was found between single leaf photosynthetic capacity and tree growth. In contrast, a high positive correlation existed between allocation to leaves and diameter growth. Results from this study suggest that allocation patterns are more affected than leaf photosynthesis in trees competing with vines.     

Do the capacity and kinetics for modification of xanthophyll cycle pool size depend on growth irradiance in temperate trees?

The present study investigated the interaction of growth irradiance (Q_int) with leaf capacity for and kinetics of adjustment of the pool size of xanthophyll cycle carotenoids (sum of violaxanthin, antheraxanthin and zeaxanthin; VAZ) and photosynthetic electron transport rate (J_max) after changes in leaf light environment. Individual leaves of lower‐canopy/lower photosynthetic capacity species Tilia cordata Mill. and upper canopy/higher photosynthetic capacity species Populus tremula L. were either illuminated by additional light of 500–800 µmol m^?2 s^?1 for 12 h photoperiod or enclosed in shade bags. The extra irradiance increased the total amount of light intercepted by two‐fold for the upper and 10–15‐fold for the lower canopy leaves, whereas the shade bags transmitted 45% of incident irradiance. In control leaves, VAZ/area, VAZ/Chl and J_max were positively associated with leaf growth irradiance (Q_int). After 11 d extra illumination, VAZ/Chl increased in all cases due to a strong reduction in foliar chlorophyll, but VAZ/area increased in the upper canopy leaves of both species, and remained constant or decreased in the lower canopy leaves of T. cordata. The slope for VAZ/area changes with cumulative extra irradiance was positively associated with Q_int only in T. cordata, but not in P. tremula. Nevertheless, all leaves of P. tremula increased VAZ/area more than the most responsive leaves of T. cordata. Shading reduced VAZ content only in P. tremula, but not in T. cordata, again demonstrating that P. tremula is a more responsive species. Compatible with the hypothesis of the role of VAZ in photoprotection, the rates of photosynthetic electron transport declined less in P. tremula than in T. cordata after the extra irradiance treatment. However, foliar chlorophyll contents of the exposed leaves declined significantly more in the upper canopy of P. tremula, which is not consistent with the suggestion that the leaves with the highest VAZ content are more resistant to photoinhibition. This study demonstrates that previous leaf light environment may significantly affect the adaptation capacity of foliage to altered light environment, and also that species differences in photosynthetic capacity and acclimation potentials importantly alter this interaction.     

Invasion impacts local species turnover in a successional system

Exotic plant invasions are often associated with declines in diversity within invaded communities. However, few studies have examined the local community dynamics underlying these impacts. Changes in species richness associated with plant invasions must occur through local changes in extinction and/or colonization rates within the community. We used long‐term, permanent plot data to evaluate the impacts of the exotic vine Lonicera japonica. Over time, species richness declined with increasing L. japonica cover. L. japonica reduced local colonization rates but had no effect on extinction rates. Furthermore, we detected significant reductions in the immigration of individual species as invasion severity increased, showing that some species are more susceptible to invasion than others. These findings suggest that declines in species richness associated with L. japonica invasion resulted from effects on local colonization rates only and not through the competitive displacement of established species.     

喷施铜和硼肥对忍冬生理生化及其茎藤指标成分的影响

以4年生忍冬为试验材料,采用盆栽和叶面喷施方式,设置不同质量浓度的铜肥(10,30,50 mg/L Cu²⁺,分别表示为Cu₁₀、Cu₃₀、Cu₅₀)和硼肥(20,40,60 mg/L B₄O^2-₇,分别表示为B₂₀、B₄₀、B₆₀)及其配施组合处理,测定忍冬叶片的光合参数以及茎藤的营养物质、药效成分含量、矿质离子含量,探究施铜和硼肥对忍冬光合生理、营养物质、矿质离子及其茎藤品质的影响,为优化忍冬栽培技术和提高其药材品质提供理论参考。结果表明：(1)喷施一定质量浓度的铜硼肥有助于提高忍冬叶片的净光合速率,尤以B₄₀Cu₅₀配施处理最佳;适宜质量浓度的铜硼肥有利于增加总叶绿素含量,高质量浓度的铜硼肥则起抑制作用。(2)配施铜硼肥对忍冬藤中营养物质及药效成分含量的影响比单施铜、硼肥更显著,其中B₄₀Cu₅₀处理可显著提高忍冬藤中可溶性蛋白、淀粉、还原糖含量,B₂₀Cu₁₀处理可显著提高可溶性糖含量;有利于忍冬藤中绿原酸和马钱苷积累的最佳配施处理分别是B₆₀Cu₅₀和B20Cu₃₀。(3)喷施铜硼肥对忍冬藤矿质离子的含量也具有显著的影响,单施与配施铜硼肥处理忍冬藤矿质离子含量均有增加,且配施铜硼肥比单施铜硼肥增加效果显著,尤以B₂₀Cu₁₀、B₂₀Cu₃₀、B₂₀Cu₅₀处理最佳。综合考虑,B₄₀Cu₅₀配施处理可有效增加忍冬藤的营养与药用指标成分含量,是忍冬叶面喷施铜和硼肥的最佳比例。     

The influence of the invasive shrub, Lonicera maackii, on leaf decomposition and microbial community dynamics

Amur honeysuckle (Lonicera maackii) is an exotic invasive shrub that is rapidly expanding into forests of eastern North America. This species forms a dense forest understory, alters tree regeneration, negatively affects herb-layer biodiversity, and alters ecosystem function. In a second-growth forest in central Kentucky, we examined the timing and production of leaf litter and compared litter chemistry, decay rates, and microbial community colonization of Amur honeysuckle to that of two native trees, white ash (Fraxinus americana) and hickory (Carya spp.). The distribution of Amur honeysuckle was clumped, allowing us to compare differences in decomposition under and away from Amur honeysuckle shrubs. Amur honeysuckle leaf litter had significantly higher nitrogen, lower C:N, and lower lignin than the other species, and decomposition rates were greater than 5×?faster. Despite the much higher rate of Amur honeysuckle decomposition compared with the native species (p?<?0.0001), decomposition of all species was significantly slower (p?=?0.0489) in sites located under Amur honeysuckle shrubs. Nitrogen concentration increased through time in decomposing Amur honeysuckle litter; however, total mass of N rapidly declined. We found the initial microbial community on leaf litter of Amur honeysuckle was distinct from two native species and although all microbial communities changed through time, the microbial community of Amur honeysuckle remained distinct from native communities. In summary, a distinct microbial community that may originate on Amur honeysuckle leaves prior to senescence could explain the rapid decay rates.     

Inferring relationships between native plant diversity and Lonicera japonica in upland forests in north Mississippi,USA

Question: Do anthropogenic disturbances interact with local environmental factors to increase the abundance and frequency of invasive species, which in turn exerts a negative effect on native biodiversity? Location: Mature Quercus‐Carya and Quercus‐Carya‐Pinus (oak‐hickory‐pine) forests in north Mississippi, USA. Methods: We used partial correlation and factor analysis to investigate relationships between native ground cover plant species richness and composition, percent cover of Lonicera japonica, and local and landscape‐level environmental variables and disturbance patterns in mature upland forests. We directly measured vegetation and environmental variables within 34 sampling subplots and quantified the amount of tree cover surrounding our plots using digital color aerial photography. Results: Simple bivariate correlations revealed that high species richness and a high proportion of herbs were associated with low Lonicera japonica cover, moist and sandy uncompacted soils, low disturbance in the surrounding landscape, and periodic prescribed burning. Partial correlations and factor analysis showed that once we accounted for the environmental factors, L japonica cover was the least important predictor of composition and among the least important predictors of species richness. Hence, much of the negative correlation between native species diversity and this invasive species was explained by soil texture and local and landscape‐level land‐use practices. Conclusions: We conclude that negative correlations between the abundance of invasive species and native plant diversity can occur in landscapes with a gradient of human disturbance, regardless of whether there is any negative effect of invasive species on native species.     

Transpiration and stomatal conductance of two wetland macrophytes (Cladium jamaicense and Typha domingensis) in the subtropical Everglades

In the subtropical Everglades, greater than 85% of water input through atmospheric precipitation, the primary source of water to the ecosystem, is lost through evapotranspiration (ET). Two dominant macrophytes that could influence ET rates through transpiration and stomatal control were investigated in a field study: Cladium jamaicense, presently the species with the greatest distribution (65%–70%); and Typha domingensis, a species increasing in distribution due to eutrophication and hydrological modifications to the ecosystem. Transpiration rates and stomatal conductance of the two species were compared at eutrophic, mesotrophic, and oligotrophic sites throughout an annual wet and dry season of subtropical Florida. Typha domingensis possessed higher transpiration and conductance rates (>; 11 mmol m^-2 sec^-1) than C. jamaicense (<;7 mmol m^-2 sec^-1; P <; 0.01), during the winter and spring months when ambient temperatures and vapor pressure were at a minimum. However, rates for the two species converged and were not significantly different during the summer and fall months. Stomatal conductance of C. jamaicense remained constant over the range of ambient vapor pressures, but significantly increased in T. domingensis with decreasing vapor pressure. Vapor pressure and light were the best predictors of seasonal and diurnal transpiration rates of T. domingensis, whereas temperature explained the most variability in C. jamaicense. Annual transpiration rates for both species were 1 to 2 mmol m^-2 sec^-1 greater at the eutrophic site than at the transitional and oligotrophic sites. Leaf area increased six times in C. jamaicense and twofold in T. domingensis from the control to nutrient-enriched sites. Results at the leaf scale suggest vegetation shifts and nutrient enrichment have the potential to alter water balances in the Everglades. However, canopy level studies may be necessary to support these conclusions when applied to the larger ecosystem or regional scale.     

Foliar production and decomposition rates in urban forests invaded by the exotic invasive shrub, <Emphasis Type="Italic">Lonicera maackii</Emphasis>

Exotic invasive shrubs can form dense monocultures in forest understories, which can have cascading effects on ecosystem structure and function. Amur honeysuckle, an exotic shrub that forms dense canopies in eastern forests, has the potential to alter plant community structure and ecosystem functions, such as primary production and decomposition. The goal of this study was to examine foliar productivity and leaf litter decomposition in forests invaded by Amur honeysuckle (Lonicera maackii) and to determine the extent to which the presence of this dominant exotic species may alter ecosystem function in these forests. We found that forests invaded by Amur honeysuckle had 16 times greater honeysuckle foliar biomass and 1.5 times lower total foliar biomass than forests of equivalent tree basal area, but having few honeysuckle shrubs. This suggests that productivity of native tree and shrub species may be reduced where honeysuckle density is high. Additionally, honeysuckle litter decayed four times faster and released nitrogen more rapidly than sugar maple litter, and sugar maple litter decayed 19% faster in forests invaded by Amur honeysuckle. These findings suggest that forests invaded by Amur honeysuckle may exhibit lower rates of organic matter accrual and less nitrogen retention in the forest floor. Since honeysuckle leaves develop in early spring before those of other shrubs or trees in the area, the rapid release of nitrogen from honeysuckle litter that we measured in early spring is timed to benefit this invasive species. The temporally coincident phenologies of nitrogen release during decomposition with the foliar growth needs of this shrub indicates that a potential positive feedback loop may exist between these processes that promotes continued growth and dominance of honeysuckle shrubs in these forested systems.     

Canopy ergodicity: can a single leaf represent an entire plant canopy?

While leaves typically emerge near shoot apices around the outer surface of a plant canopy, their relative position “moves” deeper into the canopy as additional leaves emerge. The photosynthetic capacity (A _max) of a given leaf can be expected to decline over time as its relative position (P _r) in the canopy becomes progressively deeper; this can be observed as a spatial gradient with the A _max of leaves declining distally from the shoot apex. As a consequence, we propose that the photosynthetic capacity averaged over a single leaf’s lifespan is equivalent to the average photosynthetic capacity of the entire plant canopy at a given time; in other words, there is an ergodic time to space averaging in the organization and development of plant canopies. We tested this “canopy ergodic hypothesis” in two woody (Alnus sieboldiana and Mallotus japonica) and two herbaceous (Polygonum sachalinensis and Helianthus tuberosus) species by following the photosynthetic capacity in 100 individual leaves from the time of their emergence until their death. We compared the average photosynthetic capacity of individual leaves through time (time-average) to the average photosynthetic capacity of all the leaves along a shoot at the time of emergence of the focal leaf (space-average). We found that A _max and P _r were positively correlated and that the time-averages of three plant species (Alnus, Mallotus, and Helianthus) were not significantly different from the corresponding space-averages, although the averages differed among individual plants. Polygonum, however, did show significant differences between time and space averages. Ergodicity appears to apply to the leaf–canopy relationship, at least approximately—the average photosynthetic capacity of a single leaf through time (time-average) can represent the average photosynthetic capacity of the entire canopy.     

Geometrical similarity analysis of photosynthetic light response curves, light saturation and light use efficiency

Light absorption and use efficiency (LAUE mol mol⁻¹, daily gross photosynthesis per daily incident light) of each leaf depends on several factors, including the degree of light saturation. It is often discussed that upper canopy leaves exposed to direct sunlight are fully light-saturated. However, we found that upper leaves of three temperate species, a heliophytic perennial herb Helianthus tuberosus, a pioneer tree Alnus japonica, and a late-successional tree Fagus crenata, were not fully light-saturated even under full sunlight. Geometrical analysis of the photosynthetic light response curves revealed that all the curves of the leaves from different canopy positions, as well as from the different species, can be considered as different parts of a single non-rectangular hyperbola. The analysis consistently explained how those leaves were not fully light-saturated. Light use optimization models, called big leaf models, predicted that the degree of light saturation and LAUE are both independent of light environment. From these, we hypothesized that the upper leaves should not be fully light-saturated even under direct sunlight, but instead should share the light limitation with the shaded lower-canopy leaves, so as to utilize strong sunlight efficiently. Supporting this prediction, within a canopy of H. tuberosus, both the degree of light saturation and LAUE were independent of light environment within a canopy, resulting in proportionality between the daily photosynthesis and the daily incident light among the leaves.     

A Test of Gas Exchange Measurements on Excised Canopy Branches of Ten Tropical Tree Species

We studied gas exchange of leaves on branches that had been cut and then re-cut under water to assess the utility of measuring gas exchange on leaves of excised canopy branches. There was large variation between species in their ability to photosynthesize following excision. Some species maintained up to 86.5% of intact photosynthetic rate 60 min after excision, whereas other species dropped below 40% of intact photosynthetic rates within 3 min. Three species showed significant reductions in maximum rates of gross photosynthetic rate (P _G) on leaves of excised branches relative to intact branches. Excision significantly reduced carboxylation rates (Vc _max) in four species and electron transport (J _max) in two species. There were also significant increases in compensation irradiance and reductions of day rates of respiration relative to intact measurements. While gas exchange on excised branches can provide useful measures for canopy species, responses of individual species to branch excision need to be taken into account. Measurements on pre-screened species allow a greater understanding of canopy photosynthesis of large trees when canopy access is not an option.