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.     

Partitioning of leaf nitrogen with respect to within canopy light exposure and nitrogen availability in peach (Prunus persica)

Summary Relationships between leaf nitrogen content and within canopy light exposure were studied in mature nectarine peach trees (Prunus persica cv. Fantasia) that had received 0, 112, 196, 280 or 364 kg of fertilizer nitrogen per hectare per year for the previous 3 years. The relationships between light saturated leaf CO₂ assimilation rates and leaf nitrogen concentration were also determined on trees in the highest and lowest nitrogen fertilization treatments. The slope of the linear relationship between leaf N content per unit leaf area and light exposure was similar for all nitrogen treatments but the y-intercept of the relationship increased with increasing N status. The slope of the relationship between leaf N content per unit leaf area and light saturated CO₂ assimilation rates was greater for the high N trees than the low N trees, but maximum measured leaf CO₂ assimilation rates were similar for both the high and low N treatments. A diagrammatic model of the partitioning of leaf photosynthetic capacity with respect to leaf light exposure for high and low nitrogen trees suggests that the major influence of increased N availability is an increase in the photosynthetic capacity of partially shaded leaves but not of the maximum capacity of highly exposed leaves.     

Influence of phosphorus and nitrogen on photosynthetic parameters and growth in <Emphasis Type="Italic">Vicia faba</Emphasis> L.

The influence of phosphorus (P) and nitrogen (N) supply on biomass, leaf area, photon saturated photosynthetic rate (P_max), quantum yield efficiency (α), intercellular CO₂ concentration (C_i), and carboxylation efficiency (CE) was investigated in Vicia faba. The influence of P on N accumulation, biomass, and leaf area production was also investigated. An increase in P supply was consistently associated with an increase in N accumulation and N productivity in terms of biomass and leaf area production. Furthermore, P increased the photosynthetic N use efficiency (NUE) in terms of P_max and α. An increase in P supply was also associated with an increase in CE and a decrease in C_i. Under variable daily meteorological conditions specific leaf nitrogen content (N_L), specific leaf phosphorus content (P_L), specific leaf area (δ_L), root mass fraction (R_f), P_max, and α remained constant for a given N and P supply. A monotonic decline in the steady-state value of R_f occurred with increasing N supply. δ_L increased with increasing N supply or with increasing N_L. We tested also the hypothesis that P supply positively affects both N demand and photosynthetic NUE by influencing the upper limit of the asymptotic values for P_max and CE, and the lower limit for C_i in response to increasing N.     

High resource capture and use efficiency and prolonged growth season contribute to invasiveness of Eupatorium adenophorum

To explore the traits contributing to invasion success of Eupatorium adenophorum, a noxious invasive perennial forb throughout the subtropics in Asia, Oceania, Africa, and USA, we compared the differences in ecophysiology and phenology between the invader and native E. japonicum under eight treatment combinations of two irradiances and four nitrogen additions in a two-year shadehouse experiment. The invader had significantly higher mass-based light-saturated photosynthetic rate (P _max) than its native congener in all treatments, contributing to higher photosynthetic nitrogen-, phosphorus-, and energy-use efficiencies. The higher P _max of the invader was associated with its higher nitrogen concentrations in the photosynthetic apparatus, which resulted from higher leaf nitrogen allocation to photosynthesis. The invader had higher specific leaf area and stomatal conductance at most of the treatments, also contributing to its higher P _max. The invader was not constrained by the negative correlation between leaf lifespan and specific leaf area or P _max. Leaf lifespan and total leaf area of the invader were greater than those of the native. From November to March the native congener was leafless, whereas the invader maintained a large area of leaves with relatively high P _max. Biomass accumulated in these months accounted for more than 40 % of the total biomass of the invader. Our results indicate that both the ability to capture and utilize resources efficiently and the ability to use resources when they are unavailable to natives contribute to invasion success of E. adenophorum and emphasize the importance of exploring multiple, non-mutually exclusive mechanisms for invasions.     

The Nitrogen Use Efficiency of C(3) and C(4) Plants: II. Leaf Nitrogen Effects on the Gas Exchange Characteristics of Chenopodium album (L.) and Amaranthus retroflexus (L.)

The effect of leaf nitrogen (N) on the photosynthetic capacity and the light and temperature response of photosynthesis was studied in the ecologically similar annuals Chenopodium album (C₃) and Amaranthus retroflexus (C₄). Photosynthesis was linearly dependent on leaf N per unit area (N_a) in both species. A. retroflexus exhibited a greater dependence of photosynthesis on N_a than C. album and at any given N_a, it had a greater light saturated photosynthesis rate than C. album. The difference between the species became larger as N_a increased. These results demonstrate a greater photosynthetic N use efficiency in A. retroflexus than C. album. However, at a given applied N level, C. album allocated more N to a unit of leaf area so that photosynthetic rates were similar in the two species. Leaf conductance to water vapor increased linearly with N_a in both species, but at a given photosynthetic rate, leaf conductance was higher in C. album. Thus, A. retroflexus had a greater water use efficiency than C. album. Water use efficiency was independent of leaf N in C. album, but declined with decreasing N in A. retroflexus.     

Patterns of light and nitrogen distribution in relation to whole canopy carbon gain in C3 and C4 mono- and dicotyledonous species

An analytical model was used to describe the optimal nitrogen distribution. From this model, it was hypothesized that the non-uniformity of the nitrogen distribution increases with the canopy extinction rate for light and the total amount of free nitrogen in the canopy, and that it is independent of the slope of the relation between light saturated photosynthesis (P_m) and leaf nitrogen content (n_L). These hypotheses were tested experimentally for plants with inherently different architectures and different photosynthetic modes. A garden experiment was carried out with a C₃ monocot [rice, Oryza sativa (L.)], a C₃ dicot [soybean, Glycine max (L.) Merr] a C₄ monocot [sorghum, Sorghum bicolor (L.) Moensch] and a C₄ dicot [amarantus, Amaranthus cruentus (L.)]. Leaf photosynthetic characteristics as well as light and nitrogen distribution in the canopies of dense stands of these species were measured. The dicot stands were found to have higher extinction coefficients for light than the monocot stands. Dicots also had more non-uniform N distribution patterns. The main difference between the C₃ and C₄ species was that the C₄ species were found to have a greater slope value of the leaf-level P_m—n_L relation. Patterns of N distribution were similar in stands of the C₃ and C₄ species. In general, these experimental results were in accordance with the model predictions, in that the pattern of nitrogen allocation in the canopy is mainly determined by the extinction coefficient for light and the total amount of free nitrogen.     

The Relation Between the Nitrogen Concentration and Photosynthetic Capacity of Potato (Solanum tuberosum L.) Leaves

Measurements of the rate of light-saturated photosynthesis (P_max)were made on terminal leaflets of potato plants growing in cropssupplied with 0, 3, 6, 12, 24 and 36 g N m^–2. Measurementswere made between 100 and 154 d after planting. Two types ofleaf were selected—the fourth leaf on the second-levelbranch (L₄, _B1) and the youngest terminal leaflet that was measurable(L_YM). Later, the total nitrogen concentration of each leaflet(N_L) was measured. A linear regression between P_max and N_L,common to both leaf positions, explained 68.5% of the totalvariation. With L₄, _B1 leaves there was a significant improvementin the proportion of variation explained when regressions withseparate intercepts and a common slope were fitted to individualfertilizer treatments. These results suggest that an increasingproportion of leaf nitrogen was not associated with the performanceof the photosynthetic system with increasing nitrogen supply.This separation between nitrogen treatments was not as clearfor L_YM leaves. Stomatal conductance to transfer of water vapourwas neither influenced by leaf position nor directly by nitrogensupply. Rather conductance declined in parallel with the declinein photosynthetic capacity. Solanum tuberosum, potato, nitrogen, photosynthesis, stomatal conductance, leaf     

A Cost-Benefit Analysis of Leaves of Eight Australian Savanna Tree Species of Differing Leaf Life-Span

Cost-benefit analysis of foliar construction and maintenance costs and of carbon assimilation of leaves of differing life-span were conducted using two evergreen, three semi-deciduous, and three deciduous tree species of savannas of north Australia. Rates of radiant-energy-saturated CO₂ assimilation (P _max) and dark respiration were measured and leaves were analysed for total nitrogen, fat, and ash concentrations, and for heat of combustion. Specific leaf area, and leaf N and ash contents were significantly lower in longer-lived leaves (evergreen) than shorter-lived leaves (deciduous) species. Leaves of evergreen species also had significantly higher heat of combustion and lower crude fat content than leaves of deciduous species. On a leaf area basis, P _max was highest in leaves of evergreen species, but on a leaf dry mass basis it was highest in leaves of deciduous species. P _max and total Kieldahl N content were linearly correlated across all eight species, and foliar N content was higher in leaves of deciduous than evergreen species. Leaf construction cost was significantly higher and maintenance costs were lower for leaves of evergreen than deciduous species. Maintenance and construction costs were linearly related to each other across all species. Leaves of evergreen species had a higher cost-benefit ratio compared to leaves of deciduous species but with longer lived leaves, the payback interval was longer in evergreen than deciduous species. These results support the hypotheses that longer lived leaves are more expensive to construct than short-lived leaves, and that a higher investment of N into short-lived leaves occurs which supports a higher P _max over a shorter payback interval.     

Nitrogen deposition limits photosynthetic response to elevated CO2 differentially in a dioecious species

Sexual dimorphisms of dioecious plants are important in controlling and maintaining sex ratios under changing climate environments. Yet, little is known about sex-specific responses to elevated CO₂ with soil nitrogen (N) deposition. To investigate sex-related physiological and biochemical responses to elevated CO₂ with N deposition, Populus cathayana Rehd. was employed as a model species. The cuttings were subjected to two CO₂ regimes (350 and 700???mol?mol^?1) with two N levels (0 and 5?g?N?m^?2?year^?1). Our results showed that elevated CO₂ and N deposition separately increased the total number of leaves, leaf area (LA), leaf mass, net photosynthetic rate (P _n), light saturated photosynthetic rate (P _max), chlorophyll a (Chl a), and chlorophyll a to chlorophyll b ratio (Chl a/b) in both males and females of P. cathayana. However, the effects on LA, leaf mass, P _n, P _max, Chl a and Chl a/b were weakened under the combined treatment of elevated CO₂ and N deposition. Males had higher leaf mass, P _n, P _max, apparent quantum yield (??), carboxylation efficiency (CE), Chl a, Chl a/b, leaf N, and root carbon to N ratio (C/N) than did females under elevated CO₂ with N deposition. In contrast to males, females had significantly higher levels of soluble sugars in leaves and greater starch accumulation in roots and stems under the same condition. The results of the present work imply that P. cathayana females are more responsive and suffer from greater negative effects on growth and photosynthetic capacity than do males when grown under elevated CO₂ with soil N deposition.     

The Nitrogen Use Efficiency of C(3) and C(4) Plants : III. Leaf Nitrogen Effects on the Activity of Carboxylating Enzymes in Chenopodium album (L.) and Amaranthus retroflexus (L.)

The relationships between leaf nitrogen content per unit area (N_a) and (a) the initial slope of the photosynthetic CO₂ response curve, (b) activity and amount of ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) and phosphoenolpyruvate carboxylase (PEPC), and (c) chlorophyll content were studied in the ecologically similar weeds Chenopodium album (C₃) and Amaranthus retroflexus (C₄). In both species, all parameters were linearly dependent upon leaf N_a. The dependence of the initial slope of the CO₂ response of photosynthesis on N_a was four times greater in A. retroflexus than in C. album. At equivalent leaf N_a contents, C. album had 1.5 to 2.6 times more CO₂ saturated Rubisco activity than A. retroflexus. At equal assimilation capacities, C. album had four times the Rubisco activity as A. retroflexus. In A. retroflexus, a one to one ratio between Rubisco activity and photosynthesis was observed, whereas in C. album, the CO₂ saturated Rubisco activity was three to four times the corresponding photosynthetic rate. The ratio of PEPC to Rubisco activity in A. retroflexus ranged from four at low N_a to seven at high N_a. The fraction of organic N invested in carboxylation enzymes increased with increased N_a in both species. The fraction of N invested in Rubisco ranged from 10 to 27% in C. album. In A. retroflexus, the fraction of N_a invested in Rubisco ranged from 5 to 9% and the fraction invested in PEPC ranged from 2 to 5%.     

Optimal photosynthetic use of light by tropical tree crowns achieved by adjustment of individual leaf angles and nitrogen content

Background and Aims

Theory for optimal allocation of foliar nitrogen (ONA) predicts that both nitrogen concentration and photosynthetic capacity will scale linearly with gradients of insolation within plant canopies. ONA is expected to allow plants to efficiently use both light and nitrogen. However, empirical data generally do not exhibit perfect ONA, and light-use optimization per se is little explored. The aim was to examine to what degree partitioning of nitrogen or light is optimized in the crowns of three tropical canopy tree species.

Methods

Instantaneous photosynthetic photon flux density (PPFD) incident on the adaxial surface of individual leaves was measured along vertical PPFD gradients in tree canopies at a frequency of 0·5 Hz over 9–17 d, and summed to obtain the average daily integral of PPFD for each leaf to characterize its insolation regime. Also measured were leaf N per area (N_area), leaf mass per area (LMA), the cosine of leaf inclination and the parameters of the photosynthetic light response curve [photosynthetic capacity (A_max), dark respiration (R_d), apparent quantum yield (ϕ) and curvature (θ)]. The instantaneous PPFD measurements and light response curves were used to estimate leaf daily photosynthesis (A_daily) for each leaf.

Key Results

Leaf N_area and A_max changed as a hyperbolic asymptotic function of the PPFD regime, not the linear relationship predicted by ONA. Despite this suboptimal nitrogen partitioning among leaves, A_daily did increase linearly with PPFD regime through co-ordinated adjustments in both leaf angle and physiology along canopy gradients in insolation, exhibiting a strong convergence among the three species.

Conclusions

The results suggest that canopy tree leaves in this tropical forest optimize photosynthetic use of PPFD rather than N per se. Tropical tree canopies then can be considered simple ‘big-leaves’ in which all constituent ‘small leaves’ use PPFD with the same photosynthetic efficiency.Key words: Optimal resource allocation, nitrogen, photosynthetic capacity, leaf mass per area, tropical trees, radiation use efficiency, scaling, leaf angle, canopy architecture, big leaf model     

A Comparison of Photosynthetic Capacity between Myrica Gale var. Tomentosa a Nitrogen-Fixing Shrub,and Co-Occurring Species in an Oligotrophic Moor

Photosynthetic and transpiration (E) rates, stomatal conductance, and leaf nitrogen content were surveyed for Myrica gale var. tomentosa, a N₂-fixing wetland shrub, Betula platyphylla var. japonica, and Rhododendron japonicum in Ozegahara moor, an oligotrophic moor in Central Japan. Net photosynthetic rate saturated with irradiance (P_max) of M. gale was 15.2-16.5 μmol(CO₂) m^-2 s^-1, higher than those of the other species throughout the growing season. P_max was positively correlated with leaf N content among the three species. The large leaf N content in M. gale was due to N₂-fixation in root nodules. In a comparison of M. gale in two habitats, P_max, leaf N content, and root nodule development were larger in the wetter habitat. M. gale showed high E and no midday depression of P_max even under high irradiance and large vapour pressure deficit between leaves and ambient air on a midsummer day. These traits of photosynthesis and water relations were associated with the dominance of this shrub in wetter sites such as stream sides and hollows. This revised version was published online in September 2006 with corrections to the Cover Date.     

Canopy structure and vertical patterns of photosynthesis and related leaf traits in a deciduous forest

Canopy structure and light interception were measured in an 18-m tall, closed canopy deciduous forest of sugar maple (Acer saccharum) in southwestern Wisconsin, USA, and related to leaf structural characteristics, N content, and leaf photosynthetic capacity. Light attenuation in the forest occurred primarily in the upper and middle portions of the canopy. Forest stand leaf area index (LAI) and its distribution with respect to canopy height were estimated from canopy transmittance values independently verified with a combined leaf litterfall and point-intersect method. Leaf mass, N and A _max per unit area (LMA, N/area and A _max/area, respectively) all decreased continuously by over two-fold from the upper to lower canopy, and these traits were strongly correlated with cumulative leaf area above the leaf position in the canopy. In contrast, neither N concentration nor A _max per unit mass varied significantly in relation to the vertical canopy gradient. Since leaf N concentration showed no consistent pattern with respect to canopy position, the observed vertical pattern in N/area is a direct consequence of vertical variation of LMA. N/area and LMA were strongly correlated with A _max/area among different canopy positions (r²=0.81 and r²=0.66, respectively), indicating that vertical variation in area-based photosynthetic capacity can also be attributed to variation in LMA. A model of whole-canopy photosynthesis was used to show that observed or hypothetical canopy mass distributions toward higher LMA (and hence higher N/area) in the upper portions of the canopy tended to increase integrated daily canopy photosynthesis over other LMA distribution patterns. Empirical relationships between leaf and canopy-level characteristics may help resolve problems associated with scaling gas exchange measurements made at the leaf level to the individual tree crown and forest canopy-level.     

Steady-state and dynamic photosynthetic performance and nitrogen partitioning in the shade-demanding plant Panax notoginseng under different levels of growth irradiance

In this study, we examined steady-state and dynamic photosynthetic performance and leaf nitrogen (N) partitioning in the typical shade-demanding herb Panax notoginseng grown along a light gradient. Gas exchange on a leaf area basis was significantly reduced under low irradiance, with gas exchange on a leaf mass basis reaching a maximum value and then decreasing along the light gradient. Specific leaf area significantly increased with decreasing irradiance levels (P < 0.001), whereas carboxylation efficiency was decreased (P < 0.001). In addition, decreasing growth irradiance levels led to declines in maximum carboxylation rate (V _cmax) and maximum electron transport rate (J _max), although V _cmax/mass and J _max/mass were relatively less affected than V _cmax/area and J _max/area. Slow photosynthetic response to simulated sunflecks was observed under low levels of growth irradiance, with stomatal limitations only detected in leaves grown under low-light conditions. Chlorophyll content increased significantly with decreasing irradiance levels. N content on a leaf mass basis apparently increased, while N content on a leaf area basis markedly decreased. The fraction of leaf N allocated to light-harvesting components increased significantly with decreasing growth irradiance levels, whereas the fraction allocated to carboxylation and bioenergetics was significantly reduced. As an adaptation strategy to growth irradiance, we conclude that adjustments in specific leaf area may be more important than changes in leaf physiology and biochemistry in typical shade-demanding species such as P. notoginseng.     

THE EFFECT OF GROWTH IRRADIANCE ON THE COUPLING OF CARBON AND NITROGEN METABOLISM IN CHAETOMORPHA LINUM (CHLOROPHYTA)

The influence of growth irradiance on the non-steady-state relationship between photosynthesis and tissue carbon (C) and nitrogen (N) pools in Chaetomorpha linum (Muller) Kutzing in response to abrupt changes in external nitrogen (N) availability was determined in laboratory experiments. For a given thallus N content, algae acclimated to low irradiance consistently had a higher rate of light-saturated photosynthesis (P_max normalized to dry weight) than algae acclimated to saturating irradiance; for both treatments, P_max was correlated to thallus N. Both P_max and the photosynthetic efficiency (α_dw) were correlated in C. linum grown at either saturating or limiting irradiance over the range of experimental conditions, indicating that variations in electron transport were coupled to variations in C-fixation capacity despite the large range of tissue N content from 1.1% to 4.8%. Optimizing both α and P_max and thereby acclimating to an intermediate light level may be a general characteristic of thin-structured opportunistic algae that confers a competitive advantage in estuarine environments in which both light and nutrient conditions are highly variable. Nitrogen-saturated algae had the same photosynthesis–irradiance relationship regardless of light level. When deprived of an external N supply, photosynthetic rates did not change in C. linum acclimated to low irradiance despite a two-fold decrease in tissue N content, suggesting that the active pools of chlorophyll and Rubisco remained constant. Both α and P_max decreased immediately and continuously in algae acclimated to high irradiance on removal of the N supply even though tissue N content was relatively high during most of the N-starvation period, indicating a diversion of energy and reductant away from C fixation to support high growth rates. Carbon and nitrogen assimilation were equally balanced in algae in both light treatments throughout the N-saturation and -depletion phases, except when protein synthesis was limited by the depletion of internal N reserves in severely N-starved high-light algae and excess C accumulated as starch stores. This suggests that the ability for short-term adjustment of internal allocation to acquire N andC in almost constant proportions may be especially beneficial to macroalgae living in environments characterized by high variability in light levels and nutrient supply.     

Changes in needle nitrogen partitioning and photosynthesis during 80 years of tree ontogeny in Picea abies

Needle nitrogen partitioning and photosynthesis of Norway spruce were studied in a forest chronosequence in Järvselja Experimental Forest, Estonia. Current- and previous-year shoots were sampled from upper and lower canopy positions in four stands, ranging in age from 13 to 82 years. A/c _i curves were determined to obtain maximum carboxylation rate (V _cmax) and maximum rate of electron transport (J _max), whereas needle nitrogen partitioning into carboxylation (P _R), bioenergetics associated with electron transport (P _B) and thylakoid light harvesting components (P _L) was calculated from the values of V _cmax, J _max and leaf chlorophyll concentration. The greatest changes in studied needle characteristics took place between tree ages of 13 and 26 years, and this pattern was independent of needle age and canopy position. Needle mass per projected area (LMA) was lowest in the 13-year-old stand and mass-based nitrogen concentration (N^M) was generally highest in that stand. The values of LMA were significantly higher and those of N^M lower in the 26-year-old stand. Mass-based V _cmax and J _max were highest in the 13-year-old stand. Area-based photosynthetic capacity was independent of tree age. The proportion of photosynthetic nitrogen (P _R, P _B and P _L) was highest and that of non-photosynthetic nitrogen lowest in the 13-year-old stand. Current-year needles had lower LMA and P _L, but higher photosynthetic capacity compared to 1-year-old foliage. Needles from lower canopy positions exhibited lower LMA, area-based nitrogen concentration and photosynthetic capacity than needles from upper canopy. The period of substantial reductions in needle photosynthetic capacity and changes in nitrogen partitioning coincides with the onset of reproductive phase during tree ontogeny.     

Rice varietal differences in responses of stomatal gas exchange to supplemental nitrogen application

In the field, supplemental application of N fertilizer to rice (Oryza sativa) shortly before the beginning of heading stage increases leaf N content and enhances photosynthesis during the grain-filling period. In search of varietal differences in leaf gas exchange in response to supplemental N application, we examined 13 rice varieties grown in the field during two successive years. The varieties included japonica and indica varieties, both of which are widely grown in Japan. The response to supplemental N application could not be separated clearly between variety groups; some of the japonica varieties, but none of the indica varieties, exhibited significant increase in stomatal conductance (g _s) after supplemental N application. Supplemental N was more effective to increase stomatal aperture in the varieties with inherently lower g _s. Varieties that showed greater response of g _s to supplemental N application might be able to adjust their stomatal aperture with appropriate N control. Although the internal-to-ambient CO₂ mole fraction ratio and the leaf carbon isotopic composition (δ¹³C) differed among varieties as a result of variations in stomatal aperture and the CO₂ requirement of mesophyll, supplemental N application barely influenced these parameters, because it only moderately affected stomatal aperture. Since δ¹³C tended to increase with increasing number of days from transplantation to heading stage in japonica varieties, δ¹³C values were more sensitive to differences in growth rate between years than to N application.     

Partial compensation in Psychotria marginata (Rubiaceae) after simulated defoliation increases photon capture and photosynthesis

We used Y-plant, a computer-based model of plant crown architecture analysis, to simulate effects of defoliation on daily canopy carbon gain in Psychotria marginata (Rubiaceae) plants under two contrasting irradiances. Five levels of defoliation were simulated using two different types of leaf blade damage. Compensatory increases in photon-saturated photosynthetic capacity (P _max) of 25, 50, and 100 % defoliation were also simulated. In all simulations daily photon capture and CO₂ assimilation increased with defoliation. However, without a compensatory response, daily canopy carbon gain also decreased with defoliation. Under high irradiance, reduction in daily canopy carbon gain was less than what would be expected if the response was proportional to leaf area reduction. Thus, 25 and 50 % defoliation resulted in only 20 and 41 % of daily canopy carbon gain reduction, respectively. In the scenario where 25 % of the leaf area was removed, if the P _max value was increased by 25 %, the remaining leaves compensated for 94 % of the daily canopy carbon relative to an undamaged non-compensated plant. At the same defoliation level, incrementing P _max values by 50 and 100 % resulted in overcompensation. Hence, because the increment of daily photon capture and CO₂ assimilation after defoliation was more a passive consequence of the reduction in leaf area than an active response, under the conditions tested photosynthetic compensation could be only possible through an active mechanism such as the increment of P _max values.     

Photosynthesis and photosynthate partitioning in n(2)-fixing soybeans

Leaf area, chlorophyll content, net CO₂ photoassimilation, and the partitioning of fixed carbon between leaf sucrose and starch and soluble protein were examined in Glycine max (L) Merr. cv Williams grown under three different nitrogen regimes. One group (Nod+/+) was inoculated with Bradyrhizobium and watered daily with a nutrient solution containing 6 millimolar NH₄NO₃. A second set (Nod+/−) was inoculated and had N₂ fixation as its sole source of nitrogen. A third group (Nod⁻) was not inoculated and was watered daily with a nutrient solution containing 6 millimolar NH₄NO₃. The mean net micromole CO₂ uptake per square decimeter per hour of the most recently matured source leaves was similar among the three groups of plants, being about 310. Mean leaf area of the source leaves, monitored for net photosynthesis was also similar. However, the mean milligram of chlorophyll per square decimeter of Nod+/− test leaves was about 50% lower than the other groups' leaves and indicated nitrogen deficiency. Thus, Nod+/− utilized their chlorophyll more efficiently for photosynthetic CO₂ uptake than the plants of the other treatments. The ratio of foliar carbohydrate:protein content was high in Nod+/− but low in the plants from the other two treatments. This inverse relationship between foliar protein and carbohydrate content suggests that more fixed carbon is diverted to the synthesis of protein when nitrogen availability is high. It was also found that Nod+/− sequestered more storage protein in their paraveinal mesophyll than plants of the other treatments. This study indicates that when inorganic nitrogen regimes are used to control photosynthate partitioning, then both leaf carbohydrate and leaf protein must be considered as end products of carbon assimilate allocation.     

苹果密植园与间伐园树冠层内叶片光合潜力比较

通过对成龄苹果密植园和间伐园树冠不同层次和部位叶片光合潜力及辐射通量密度、叶片N含量和比叶重等指标的比较分析,研究了苹果园改造前后辐射能和氮素利用效率差异及其与产量品质的关系.结果表明：间伐显著改善了冠层内的辐射环境,间伐园冠层内的辐射分布明显比密植园均匀,相对辐射通量密度小于30%的无效光区接近0,而密植园冠层内的最低相对辐射通量密度为17%,在相对高度03以下均为无效光区;间伐园内冠层叶片的光合效率显著提高,间伐园树冠中、下部叶片的光合速率比密植园分别提高了78%和102%;叶片的最大羧化速率和最大电子传递速率也有较大幅度的提升.苹果园冠层叶片的光合效率与叶片N含量存在显著的相关关系,而叶片N含量又与辐射通量密度存在显著的相关关系,因此,可根据冠层叶片相对N含量的垂直分布间接和定量地判断叶片的光合效率或相对辐射通量密度的空间分布.