Shimpei Oikawa

Does leaf shedding increase the whole-plant carbon gain despite some nitrogen being lost with shedding?

When old leaves are shed, part of the nitrogen in the leaf is retranslocated to new leaves. This retranslocation will increase the whole-plant carbon gain when daily C gain : leaf N ratio (daily photosynthetic N-use efficiency, NUE) in the old leaf, expressed as a fraction of NUE in the new leaf, becomes lower than the fraction of leaf N that is resorbed before shedding (R(N)). We examined whether plants shed their leaves to increase the whole-plant C gain in accord with this criterion in a dense stand of an annual herb, Xanthium canadense, grown under high (HN) and low (LN) nitrogen availability. The NUE of a leaf at shedding expressed as a fraction of NUE in a new leaf was nearly equal to the R(N) in the LN stand, but significantly lower than the R(N) in the HN stand. Thus shedding of old leaves occurred as expected in the LN stand, whereas in the HN stand, shedding occurred later than expected. Sensitivity analyses showed that the decline in NUE of a leaf resulted primarily from a reduction in irradiance in the HN stand. On the other hand, it resulted from a reduction in irradiance and also in light-saturated photosynthesis : leaf N content ratio (potential photosynthetic NUE) in the LN stand.

Effects of elevated CO2 concentration on seed production in C3 annual plants

The response of seed production to CO(2) concentration ([CO(2)]) is known to vary considerably among C(3) annual species. Here we analyse the interspecific variation in CO(2) responses of seed production per plant with particular attention to nitrogen use. Provided that seed production is limited by nitrogen availability, an increase in seed mass per plant results from increase in seed nitrogen per plant and/or from decrease in seed nitrogen concentration ([N]). Meta-analysis reveals that the increase in seed mass per plant under elevated [CO(2)] is mainly due to increase in seed nitrogen per plant rather than seed [N] dilution. Nitrogen-fixing legumes enhanced nitrogen acquisition more than non-nitrogen-fixers, resulting in a large increase in seed mass per plant. In Poaceae, an increase in seed mass per plant was also caused by a decrease in seed [N]. Greater carbon allocation to albumen (endosperm and/or perisperm) than the embryo may account for [N] reduction in grass seeds. These differences in CO(2) response of seed production among functional groups may affect their fitness, leading to changes in species composition in the future high-[CO(2)] ecosystem.

Light interception in species with different functional groups coexisting in moorland plant communities

Competition for light is one of the most essential mechanisms affecting species composition. It has been suggested that similar light acquisition efficiency (Φmass, absorbed photon flux per unit aboveground mass) may contribute to species coexistence in multi-species communities. On the other hand, it is known that traits related with light acquisition vary among functional groups. We studied whether Φmass was similar among species with different functional groups coexisting in moorland communities. We conducted stratified clipping in midsummer when the stand biomass reached a maximum. Light partitioning among species was estimated using a model accounting for both direct and diffuse light. Evergreen species were found to have a significantly lower Φmass than deciduous species, which resulted from their lower absorbed photon flux per unit leaf area and lower specific leaf area. Shrubs had a smaller leaf mass fraction, but their Φmass was not lower than that of herbs because they had a higher leaf position due to the presence of wintering stems. Species with vertical leaves had a higher Φmass than those with horizontal leaves despite vertical leaves being a decided disadvantage in terms of light absorption. This higher Φmass was achieved by a greater leaf height in species with vertical leaves. Our results clearly demonstrate that light acquisition efficiency was different among the functional groups. However, the trend observed is not necessarily the same as that expected based on prior knowledge, suggesting that disadvantages in some traits for light acquisition efficiency are partly compensated for by other traits.

Dynamics of leaf area and nitrogen in the canopy of an annual herb, Xanthium canadense

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.

Demand and supply of N in seed production of soybean (Glycine max) at different N fertilization levels after flowering

Nitrogen (N) has been suggested as a determinant of seed production especially in species with high seed N content. Assuming that seed yield was determined as the balance between N demand and supply for seed production, we studied the effect of N fertilization after flowering on soybean (Glycine max L. Merr.) yield. Seed N concentration was nearly constant irrespective of N fertilization, indicating that seed production was proportional to the amount of N available for seed growth. N demand for seed production was analyzed as the product of seed number, the rate of N filling in individual seeds, and the length of the reproductive period. N fertilization increased seed number and the reproductive period, but did not influence the N filling rate. Seed number was positively correlated with dry mass productivity after flowering. Three N sources were distinguished: mineral N uptake, symbiotic N2 fixation and N remobilization from vegetative body. N fertilization increased N uptake and N remobilization, but lowered N2 fixation. We concluded that N availability in the reproductive period determined seed yield directly through increasing N supply for seed growth and indirectly through increasing seed N demand with enhanced plant dry mass productivity.

A meta-analysis of leaf nitrogen distribution within plant canopies

BACKGROUND AND AIMS Leaf nitrogen distribution in the plant canopy is an important determinant for canopy photosynthesis. Although the gradient of leaf nitrogen is formed along light gradients in the canopy, its quantitative variations among species and environmental responses remain unknown. Here, we conducted a global meta-analysis of leaf nitrogen distribution in plant canopies. METHODS We collected data on the nitrogen distribution and environmental variables from 393 plant canopies (100, 241 and 52 canopies for wheat, other herbaceous and woody species, respectively). KEY RESULTS The trends were clearly different between wheat and other species; the photosynthetic nitrogen distribution coefficient (Kb) was mainly determined by leaf area index (LAI) in wheat, whereas it was correlated with the light extinction coefficient (KL) and LAI in other species. Some other variables were also found to influence Kb We present the best equations for Kb as a function of environmental variables and canopy characteristics. As a more simple function, Kb = 0·5KL can be used for canopies of species other than wheat. Sensitivity analyses using a terrestrial carbon flux model showed that gross primary production tended to be more sensitive to the Kb value especially when nitrogen content of the uppermost leaf was fixed. CONCLUSION Our results reveal that nitrogen distribution is mainly driven by the vertical light gradient but other factors such as LAI also have significant effects. Our equations contribute to an improvement in the projection of plant productivity and cycling of carbon and nitrogen in terrestrial ecosystems.

Mean residence time of leaf number, area, mass, and nitrogen in canopy photosynthesis

Mean residence time (MRT) of plant nitrogen (N), which is an indicator of the expected length of time N newly taken up is retained before being lost, is an important component in plant nitrogen use. Here we extend the concept MRT to cover such variables as leaf number, leaf area, leaf dry mass, and nitrogen in the canopy. MRT was calculated from leaf duration (i.e., time integral of standing amount) divided by the total production of leaf variables. We determined MRT in a Xanthium canadense stand established with high or low N availability. The MRT of leaf number may imply longevity of leaves in the canopy. We found that the MRT of leaf area and dry mass were shorter than that of leaf number, while the MRT of leaf N was longer. The relatively longer MRT of leaf N was due to N resorption before leaf shedding. The MRT of all variables was longer at low N availability. Leaf productivity is the rate of canopy photosynthesis per unit amount of leaf variables, and multiplication of leaf productivity by MRT gives the leaf photosynthetic efficiency (canopy photosynthesis per unit production of leaf variables). The photosynthetic efficiency of leaf number implies the lifetime carbon gain of a leaf in the canopy. The analysis of plant-level N use efficiency by evaluating the N productivity and MRT is a well-established approach. Extension of these concepts to leaf number, area, mass, and N in the canopy will clarify the underlying logic in the study of leaf life span, leaf area development, and dry mass and N use in canopy photosynthesis.

Effects of elevated CO2 on leaf area dynamics in nodulating and non-nodulating soybean stands

AimsThe effects of elevated CO2 on leaf area index (LAI) vary among studies. We hypothesized that the interactive effects of CO2 and nitrogen on leaf area loss have important roles in LAI regulation.MethodsWe studied the leaf area production and loss using nodulating soybean and its non-nodulating isogenic line in CO2-controlled greenhouse systems.ResultsLeaf area production increased with elevated CO2 levels in the nodulating soybean stand and to a lesser extent in the non-nodulating line. Elevated CO2 levels accelerated leaf area loss only in nodulating plants. Consequently, both plants exhibited a similar stimulation of peak LAI with CO2 elevation. The accelerated leaf loss in nodulating plants may have been caused by newly produced leaves shading the lower leaves. The nodulating plants acquired N throughout the growth phase, whereas non-nodulating plants did not acquire N after flowering due to the depletion of soil N. N retranslocation to new organs and subsequent leaf loss were faster in non-nodulating plants compared with nodulating plants, irrespective of the CO2 levels.ConclusionLAI regulation in soybean involved various factors, such as light availability within the canopy, N acquisition and N demands in new organs. These effects varied among the growth stages and CO2 levels.

Nitrogen-utilization efficiency in rice: an analysis at leaf, shoot, and whole-plant level

AimsIf rice has a higher nitrogen use efficiency for dry matter production (NUE) and grain yield (NUEGY) than wild annuals, we may question whether the higher NUE is due to a higher productivity per plant N (NP) or a longer retention time of plant N (MRT) or both, and whether the higher NUEGY results also from a higher harvest index (HI).MethodsStands of rice were established at three N levels. Censuses were done for birth and death of every shoot and leaf from germination to full maturity. Nitrogen uptake, dry matter production, grain yield, HI, NUE, and NUEGY were determined at shoot and whole-plant levels.ResultsRice had a higher NUE, NUEGY and NP, but hardly higher plant-N MRT and HI than wild annuals. Leaf-N MRT was higher than leaf longevity in fertile shoots, while the opposite was true in sterile tillers. Shoot NUEGY was higher in late tillers due to higher HI.ConclusionsHigh NUE of rice results from its high NP, not from high MRT at both whole-plant and leaf levels. Revenues gained from enhancing MRT would have been lower than the opportunity costs of reducing NP. N recycles between shoots. Sterile tillers function as an N storage for grain yield in fertile shoots.

Seasonal change in light partitioning among coexisting species of different functional groups along elevation gradient in subalpine moorlands

Species niches are expected to differ between different functional groups and between species with different functional traits. However, it is still unclear how functional traits contribute to niche separation between species coexisting in a community and between sites along environmental gradients. We studied seasonal changes in light partitioning among coexisting species belonging to different functional groups in moorland plant communities at different altitudes. We estimated the lifetime light absorption per unit invested leaf biomass (ΦLleafmass) as a measure of the benefit/cost ratio of light acquisition. Evergreen species absorbed more light in spring, whereas deciduous species absorbed more light in summer. A similar tradeoff was also found between short and tall species within each functional group. As a result, evergreen and shorter species had comparable ΦLleafmass values to those of deciduous and taller species. Evergreen species had higher ΦLleafmass at higher altitudes relative to deciduous species, suggesting that evergreen habit is more advantageous for the lifetime light interception at higher altitudes. Our results demonstrate that phenological tradeoffs for light partitioning can contribute to the coexistence of species with different functional traits. Our results also reveal that the most advantageous traits differ depending on environment.