Effects of different concentrations of nitrogen and phosphorus on chlorophyll biosynthesis, chlorophyll a fluorescence, and photosynthesis in Larix olgensis seedlings

In our experiments, one-year-old Larix olgensis seedlings were cultivated in sand, and supplied with solutions with different concentrations of nitrate or phosphate. The effects of nitrogen and phosphorus supply on chlorophyll biosynthesis, total nitrogen content, and photosynthetic rate were studied. The experimental results are listed below: 1) 5-aminolevulinic acid (ALA) synthetic rate increased as nitrate concentrations supplied to larch seedlings increased from 1 to 8 mmol/L. But the rate decreased by 17% when nitrate concentration increased to 16 mmol/L, in contrast to the control. Under phosphate treatments, ALA synthetic rates were similar to those under nitrate treatments. The activities of porphobilinogen (PBG) synthase reached a maximum when larch seedlings were supplied with 8 mmol/L of nitrate or 1 mmol/L of phosphate. 2) when larch seedlings were supplied with 8 mmol/L of nitrate and 0.5 mmol/L of phosphate, the contents of chlorophyll a, chlorophyll b, total chlorophyll, and carotenoids reached a maximum. The total nitrogen contents in leaves increased as nitrate concentrations increased. 3) When phosphate concentrations increased from 0.125 to 1 mmol/L, the total nitrogen contents in leaves slightly increased; however, continuous increase of phosphate concentrations resulted in the decrease in total nitrogen contents in leaves. When nitrate concentrations increased from 1 to 8 mmol/L, soluble protein contents in leaves increased in general, and continuous increase of nitrate concentrations induced a decrease in soluble protein contents in leaves. Under treatment of 0.25 mmol/L of phosphate, the soluble protein contents reached a maximum. 4) In general, F _v/F _m increased as nitrate concentrations increased from 1 to 8 mmol/L, and continuous increase of nitrate concentration resulted in decrease in F _v/F _m. The similar changes occurred under phosphate treatments. As nitrate concentrations increased from 1 to 8 mmol/L, photosynthetic rates gradually increased, but when nitrate concentrations increased to 16 mmol/L, photosynthetic rate reduced by 16%, in contrast to the control. Photosynthetic rates reached a maximum when seedlings were supplied with 1 mmol/L, and an oversupply of phosphate (2 mmol/L) resulted in decrease in photosynthetic rates. The results suggested that supply levels of nitrogen affected ALA biosynthetic rates, activities of PBG synthase, and affected contents of chlorophyll and carotenoids. Moreover, nitrogen supply levels affected contents of total nitrogen and soluble proteins in leaves, and net photosynthetic rates. ALA biosynthesis rates and activities of PBG synthase were affected by phosphate supply, but contents of chlorophyll and carotenoids were not affected. And net photosynthetic rates were affected little by phosphate supply. __________ Translated from Scientia Silvae Sinicae, 2005, 41(4) [译自:林业科学, 2005, 41(4)]     

Stock-type patterns of phosphorus uptake,retranslocation, net photosynthesis and morphological development in interior spruce seedlings

One-year-old interior spruce (Picea glauca (Moench) Voss × Picea engelmannii Parry) spring-stock and summer-stock were grown under two phosphorus (P) fertility regimes, with (+P) or without (–P), followed by a simulated winter, and a second growing period under an adequate fertility regime in a controlled environment room. The two stock-types differed in their response to low P availability. For spring-stock, morphological development, phosphorus-use efficiency (PUE) and P specific absorption rate (SAR) were similar between –P and +P seedlings. For summer-stock, –P seedlings compared to +P seedlings had lower (p 0.05) morphological development, but greater PUE and SAR. For both stock-types, P content increased in +P seedlings, remained low in –P seedlings, and P concentration decreased in nursery-needles (i.e., formed in the nursery) of –P seedlings. The difference in stock-type response to low phosphorus availability (–P) was attributed to internal supply of P and it's retranslocation. Assimilation (A) of CO₂ in nursery-needles was similar between –P and +P seedlings for both stock-types. For spring-stock, +P seedlings had greater A in new-needles (i.e., needles formed during the trial) than –P seedlings. It was recommended that the spring-stock be selected over summer-stock for sites low in P availability.     

Morphological and physiological parameters influence the use efficiency of nitrogen and phosphorus by Eucalyptus seedlings

New Forests - Morpho-physiological characteristics in Eucalyptus species can determine N (NUE) and P (PUE) use efficiency and, consequently, the plant's responses to fertilization. The study...     

Leaf growth, photosynthesis and tissue water relations of greenhouse-grown Eucalyptus marginata seedlings in response to water deficits

Leaf growth, rate of leaf photosynthesis and tissue water relations of shoots of Eucalyptus marginata Donn ex Sm. (jarrah) seedlings were studied during a soil drying and rewatering cycle in a greenhouse experiment. Rates of leaf growth and photosynthesis were sensitive to water deficits. The rate of leaf growth decreased linearly with predawn leaf water potential to reach zero at -1.5 MPa. Rate of leaf growth did not recover completely within the first three days after rewatering. Midday photosynthetic rates declined to 40% of those of well-watered seedlings at a predawn leaf water potential of -1.0 MPa and reached zero at -2.2 MPa. Photosynthetic rate recovered rapidly following rewatering and almost fully recovered by the second day after rewatering. All tissue water relations parameters, except the bulk modulus of elasticity, changed significantly as the soil dried and recovered completely by the third day after rewatering. Changes in osmotic pressure at full turgor of 0.4 MPa indicated considerable capacity for osmotic adjustment. However, because there was little osmotic adjustment until predawn leaf water potential fell below -1.5 MPa, this capacity would not have enhanced seedling growth, although it may have increased seedling survival. The sensitivity of photosynthesis and relative water content to water deficits suggests that greenhouse-grown E. marginata seedlings behave like mesophytic plants, even though E. marginata seedlings naturally grow in a drought-prone environment.     

氮磷配比对水曲柳光合作用的影响

以水曲柳2年生苗木作为试验材料,在人工气候室中用霍格兰水培营养液修改配方(四种氮、磷的不同配比,清水作为对照)进行水培试验,测定了叶片的光合作用、叶绿素荧光以及叶绿素含量等,以期从光合作用角度探讨不同配比的氮、磷对水曲柳幼苗生产力的影响。结果表明:不同氮、磷配比对水曲柳的多个光合指标差异影响显著。在一定范围内随着施磷量的逐渐增加净光合速率(Pn)升高;施磷肥对提高水曲柳叶片光合作用能力有积极作用;磷胁迫使叶绿素含量降低且对水曲柳叶片的碳同化和光能利用产生负影响。     

Increased photosynthesis following partial defoliation of field-grown Eucalyptus globulus seedlings is not caused by increased leaf nitrogen

Increased photosynthetic rates following partial defoliation may arise from changes in leaf biochemistry, water relations or nutrient status. Twelve-month-old field-grown Eucalyptus globulus Labill. seedlings were pruned from below to reduce the green crown depth by 50 (D50) or 70% (D70). Photosynthetic responses to light and CO2 concentration were examined before and one, three and five weeks after partial defoliation. One week after defoliation, photosynthetic rates were greater in seedlings in the D50 (21 micromol m(-2) s(-1)) and D70 (23 micromol m(-2) s(-1)) treatments than in control seedlings (15 micromol m(-2) s(-1)); however, there was little difference in photosynthetic rates between partially defoliated seedlings and control seedlings after 5 weeks. An analysis of the sensitivity of photosynthesis to biochemical parameters revealed that the transient increase in photosynthetic rate in response to partial defoliation was largely a function of the maximum rate of carboxylation (85-87%) and the maximum rate of RuBP regeneration (55-60%) rather than stomatal conductance (12-13%). Nitrogen increased in leaves following partial defoliation (increases of 0.6 and 1.2 g m(-2) for D50 and D70, respectively), but was accumulated in a non-photosynthetic form (i.e., there was no increase in nitrogen concentration of Rubisco or chlorophyll). Increased photosynthetic rates immediately following partial defoliation were primarily a result of increased activity rather than amount of photosynthetic machinery. There was no evidence that phosphorus was responsible for the increase in photosynthetic rates after partial defoliation.     

Leaf growth of Eucalyptus globulus seedlings under water deficit

Eucalyptus globulus Labill. seedlings grown under field conditions in Portugal were watered either daily (control) or every 6 days (drought-treated). Relative to those of control plants, rates of leaf production and leaf biomass accumulation were reduced by almost half in drought-treated plants. However, whereas expansion of new leaves on control plants slowed toward the end of the 30 day experiment, expansion of leaves of the same age on drought-treated plants accelerated as a change in weather conditions resulted in midday plant water potentials above -3.0 MPa. In plants that were left unwatered until they wilted and were then watered daily, expansion of the fifth leaf pair from the apex was slower than that of the same pair of leaves of plants watered daily throughout; but it continued for about twice as long and resulted in the same final leaf area. Drought treatment also caused a substantial reduction in the rate of leaf production, which, in part, accounted for the effect of drought on leaf biomass production. In a greenhouse study, witholding water for 15 days had only a slight effect on the length or width of adaxial epidermal cells, and the effect was quickly reversed on rewatering.     

Defoliation and nitrogen effects on photosynthesis and growth of Eucalyptus globulus

Plant responses to defoliation are complex. We established a field experiment in a nine-month-old Eucalyptus globulus Labill. plantation to examine the effects of pattern (upper crown versus lower crown removal), frequency (single, double or triple defoliation within a 12-month period) and severity (25 versus 38% of leaf area removed) of defoliation and the effect of soil nitrogen (N) on photosynthetic processes and stem growth. The photosynthetic responses observed following defoliation could be attributed to changes in source:sink ratios. Light-saturated CO(2) uptake (A(max)) increased with increasing severity and frequency of defoliation irrespective of defoliation pattern. Seedlings defoliated in autumn did not exhibit increases in A(max) until the following spring, whereas there was no such delay in photosynthetic responses associated with spring defoliation. Application of N before defoliation allowed trees to compensate for the effect of defoliation on stem diameter growth, which could not be explained simply in terms of increases in A(max). The observed increases in stem diameter increment following N fertilization of defoliated trees suggested increases in leaf area development, and there were changes in the leaf area:leaf dry mass ratio that may have increased light absorption by the crown. Nitrogen fertilization also increased partitioning of dry mass to branches at the expense of main stems, suggesting that N supply was important in rebuilding crowns following a defoliation event.     

Relationships between net photosynthesis and foliar nitrogen concentrations in a loblolly pine forest ecosystem grown in elevated atmospheric carbon dioxide

We examined the effects of elevated carbon dioxide concentration ([CO2]) on the relationship between light-saturated net photosynthesis (A(sat)) and area-based foliar nitrogen (N) concentration (N(a)) in the canopy of the Duke Forest FACE experiment. Measurements of A(sat) and N(a) were made on two tree species growing in the forest overstory and four tree species growing in the forest understory, in ambient and elevated [CO2] FACE rings, during early and late summer of 1999, 2001 and 2002, corresponding to years three, five and six of CO2 treatment. When measured at the growth [CO2], net photosynthetic rates of each species examined in the forest overstory and understory were stimulated by elevated [CO2] at each measurement date. We found no effect of elevated [CO2] on N(a) in any of the species. The slope of the A(sat)-N relationship was 81% greater in elevated [CO2] than in ambient [CO2] when averaged across all sample dates, reflecting a differential CO2 effect on photosynthesis at the top and bottom of the canopy. We compared A(sat)-N relationships in trees grown in ambient and elevated [CO2] at two common CO2 concentrations, during late summer 2001 and both early and late 2002, to determine if the stimulatory effect of elevated [CO2] on photosynthesis diminishes over time. At all three sample times, neither the slopes nor the y-intercepts of the A(sat)-N relationships of trees grown in ambient or elevated [CO2] differed when measured at common CO2 concentrations, indicating that the responses of photosynthesis to long-term elevated [CO2] did not differ from the responses to a short-term increase in [CO2]. This finding, together with the observation that N(a) was unaffected by growth in elevated [CO2], indicates that these overstory and understory trees growing at the Duke Forest FACE experiment continue to show a strong stimulation of photosynthesis by elevated [CO2].     

Cyanogenesis in Eucalyptus polyanthemos seedlings: heritability, ontogeny and effect of soil nitrogen

Cyanogenic plants release cyanide from endogenous cyanide-containing compounds (generally cyanogenic glycosides) and thus have an effective means of chemical defense. The capacity for cyanogenesis can be highly variable, even among individuals within a population. The genetic, environmental and developmental factors determining this variability are poorly understood, particularly in tree species. We used Eucalyptus polyanthemos Schauer subsp. vestita L. Johnson & K. Hill to quantify aspects of the regulation of cyanogenic capacity, which in this species is determined by foliar cyanogenic glycoside concentration. A half-sibling progeny trial, based on seed collected from open-pollinated trees covering a range of cyanogenic capacities, was used to assess the heritability of cyanogenesis in E. polyanthemos. Narrow sense heritability (h(2) +/- 1 SE) was estimated to be 0.82 +/- 0.20 from an intra-class correlation and 0.78 +/- 0.11 from a standardized progeny-parent regression. Foliar cyanogenic glycoside concentrations were on average about 70% lower in seedlings than in maternal trees, suggesting that there is a developmental delay in the accumulation of cyanogenic capacity in this species. The high h(2) values indicate that cyanogenic capacity is largely genetically determined and that environmental factors have little effect. To test this supposition, we grew seedlings at two soil nitrogen (N) concentrations (N influences cyanogenic capacity in some species) and found no appreciable effect on cyanogenic glycoside concentration, biomass partitioning or relative growth rate. Highly cyanogenic seedlings grew more slowly than seedlings with lower cyanogenic capacities, and relative growth rate was positively associated with net assimilation rate in seedlings in both N treatments.     

Leaf senescence and late-season net photosynthesis of sun and shade leaves of overstory sweetgum (Liquidambar styraciflua) grown in elevated and ambient carbon dioxide concentrations

We examined the effects of elevated CO2 concentration ([CO2]) on leaf demography, late-season photosynthesis and leaf N resorption of overstory sweetgum (Liquidambar styraciflua L.) trees in the Duke Forest Free Air CO2 Enrichment (FACE) experiment. Sun and shade leaves were subdivided into early leaves (formed in the overwintering bud) and late leaves (formed during the growing season). Overall, we found that leaf-level net photosynthetic rates were enhanced by atmospheric CO2 enrichment throughout the season until early November; however, sun leaves showed a greater response to atmospheric CO2 enrichment than shade leaves. Elevated [CO2] did not affect leaf longevity, emergence date or abscission date of sun leaves or shade leaves. Leaf number and leaf area per shoot were unaffected by CO2 treatment. A simple shoot photosynthesis model indicated that elevated [CO2] stimulated photosynthesis by 60% in sun shoots, but by only 3% in shade shoots. Whole-shoot photosynthetic rate was more than 12 times greater in sun shoots than in shade shoots. In senescent leaves, elevated [CO2] did not affect residual leaf nitrogen, and nitrogen resorption was largely unaffected by atmospheric CO2 enrichment, except for a small decrease in shade leaves. Overall, elevated [CO2] had little effect on the number of leaves per shoot at any time during the season and, therefore, did not change seasonal carbon gain by extending or shortening the growing season. Stimulation of carbon gain by atmospheric CO2 enrichment in sweetgum trees growing in the Duke Forest FACE experiment was the result of a strong stimulation of photosynthesis throughout the growing season.     

Leaf and litter nitrogen and phosphorus in three forests with low P supply

We compared the N and P contents of the main labile components of nutrient cycles in three different forest ecosystems [a tropical evergreen forest (TEF); a tropical dry forest (TDF); and a Mediterranean temperate forest (MTF)] with low P supply. A mass-balance approach was used to estimate mean residence times for organic matter, N and P in the forest floor, and to examine the flexibility of N and P intra-system cycling in the three forest ecosystems. For this purpose, we combined published values of N and P in foliage, litterfall, forest floor litter and mineral soils in these three forest ecosystems. The results of our analysis were consistent with the widely held belief that the N content of leaves (both green and senescent) and litter increases with increasing temperatures. In contrast, the data did not support the hypothesis that leaf P content decreases with increasing temperatures and precipitation: leaf and litterfall P contents were higher in both tropical forests than they were in the temperate forest. The TEF had the highest P content of the three forests studied. The mass-balance analysis indicated that although P mineralization in the TDF can run ahead of litter decomposition stoichiometry when P is in short supply, flexibility is much reduced or absent in the TEF and the MTF. Our analysis provides additional evidence of the importance of climatic factors in forest ecosystem processes and highlights the role of flexibility in ecosystem nutrient cycling, especially for P in ecosystems with a limited P supply.     

Temperature effects on wood anatomy, wood density, photosynthesis and biomass partitioning of Eucalyptus grandis seedlings

Wood density, a gross measure of wood mass relative to wood volume, is important in our understanding of stem volume growth, carbon sequestration and leaf water supply. Disproportionate changes in the ratio of wood mass to volume may occur at the level of the whole stem or the individual cell. In general, there is a positive relationship between temperature and wood density of eucalypts, although this relationship has broken down in recent years with wood density decreasing as global temperatures have risen. To determine the anatomical causes of the effects of temperature on wood density, Eucalyptus grandis W. Hill ex Maiden seedlings were grown in controlled-environment cabinets at constant temperatures from 10 to 35 degrees C. The 20% increase in wood density of E. grandis seedlings grown at the higher temperatures was variously related to a 40% reduction in lumen area of xylem vessels, a 10% reduction in the lumen area of fiber cells and a 10% increase in fiber cell wall thickness. The changes in cell wall characteristics could be considered analogous to changes in carbon supply. Lumen area of fiber cells declined because of reduced fiber cell expansion and increased fiber cell wall thickening. Fiber cell wall thickness was positively related to canopy CO2 assimilation rate (Ac), which increased 26-fold because of a 24-fold increase in leaf area and a doubling in leaf CO2 assimilation rate from minima at 10 and 35 degrees C to maxima at 25 and 30 degrees C. Increased Ac increased seedling volume, biomass and wood density; but increased wood density was also related to a shift in partitioning of seedling biomass from roots to stems as temperature increased.     

Response of Ulmus americana seedlings to varying nitrogen and water status. 2 Water and nitrogen use efficiency in photosynthesis

Photosynthetic utilization of water and nitrogen in Ulmus americana L. seedlings was tightly linked with the relative availability of each resource. During periodic drying cycles, water use efficiency increased as predawn water potential fell from -0.5 to -2.0 MPa. During the later stages of such drying cycles, the relative contribution of stomatal limitations to the total net photosynthetic limitation appeared to be at its greatest, whereas biochemical limitations were predominant in well-watered plants grown under low nitrogen (N) availability. For any level of leaf water status, water use efficiency of photosynthesis (WUE) was always greater in plants with high leaf N content than in plants with low leaf N content. Photosynthetic nitrogen use efficiency (PNUE) was always greater in plants with low leaf N content than in plants with high leaf N content, for any level of water status. In combined N treatments and predawn water status classes, there was a significant inverse relationship between PNUE and WUE.     

Increase of nitrogen to promote growth of poplar seedlings and enhance photosynthesis under NaCl stress

The solution culture method was used to study the effect of increasing nitrogen on the growth and pho-tosynthesis of poplar seedlings under 100 mmol L-1 NaCl stress. I Increase in nitrogen reduced stomatal limitation of leaves under NaCl stress, improved utilization of CO2 by mesophyll cells, enhanced photosynthetic carbon assimi-lation capacity, significantly alleviated saline damage of NaCl, and promoted the accumulation of aboveground and root biomass. I Increased nitrogen enhanced photochemical efficiency (ΦPSⅡ) and electron transport rates, relieved the reduction of maximum photochemical efficiency (Fv/Fm) under NaCl, and reduced the degree of photoinhibition caused by NaCl stress. Increased nitrogen applications reduced the proportion of energy dissipating in the form of ineffective heat energy and hence a greater proportion of light energy absorbed by leaves was allocated to photo-chemical reactions. Under treatment with increased nitro-gen, the synergistic effect of heat dissipation and the xanthophyll cycle in the leaves effectively protected pho-tosynthetic PSⅡ and enhanced light energy utilization of leaves under NaCl stress. The increased nitrogen promoted photosynthetic electron supply and transport ability under NaCl stress evident in enhanced functioning of the oxygen-evolving complex on the electron donor side of PS Ⅱ. It increased the ability of the receptor pool to accept electrons on the PSII electron acceptor side and improved the sta-bility of thylakoid membranes under NaCl stress. Therefore, increasing nitrogen applications under NaCl stress can promote poplar growth by improving the effi-ciency of light energy utilization.     

Leaf chlorophyll,net gas exchange and chloroplast ultrastructure in citrus leaves of different nitrogen status

One-year-old 'Cleopatra mandarin' (Citrus reticulata Blanco) seedlings were raised in a greenhouse and fertilized with nitrogen (N) at four application frequencies. Nitrogen-deficient leaves (86 mmol N m-2) had less chlorophyll per unit area, but a greater chlorophyll a:b ratio than N-fertilized leaves (> 187 mmol N m-2). Leaf dry mass per area (DM area-1) and total chlorophyll concentration increased linearly with increasing leaf N, whereas chlorophyll a:b ratio declined. Net assimilation of CO2 (A(CO2)) and leaf water-use efficiency (WUE) reached maximum values in leaves with approximately 187 mmol N m-2. Nitrogen-deficient leaves exhibited small chloroplasts with no starch granules; grana and stroma lamellae that coincided with the accretion of numerous large plastoglobuli in the stroma disintegrated. High-N leaves had large chloroplasts with well-developed grana, stroma lamellae and starch granules that enlarged with increasing N concentration. The lack of an increase in A(CO2) capacity at leaf N concentrations above 187 mmol N m-2 appeared to be correlated with the presence of numerous large starch granules.     

Role of nitrogen remobilization from old leaves for new leaf growth of Eucalyptus globulus seedlings

Six-month-old Eucalyptus globulus Labill. seedlings were grown in sand culture irrigated with a nutrient solution containing 6.0 mol N m(-3) for 3 months (November-January). Before rapid growth began in February, seedlings were repotted and irrigated with either 6.0 mol N m(-3) (High-N treatment) or 1.0 mol N m(-3) (Low-N treatment). Seedlings were analyzed during the subsequent flush of growth to determine the role of old leaves, and in particular the leaf protein Rubsico, as a source of N for new leaf growth. During spring growth, the N content of old leaves of High-N seedlings decreased with decreasing leaf dry weight, although there was no change in leaf number. In High-N seedlings, the net loss of N from old leaves provided less than 10% of the N used for new leaf growth, and the new leaves quickly became the dominant sink for N. In contrast, in Low-N seedlings, the net loss of N from old leaves provided 44% of the N used for new leaf growth. During the period of spring growth, the amount of soluble proteins recovered from old leaves of Low-N seedlings dropped, but there was no change in the content of either Rubisco or chlorophyll. The photosynthetic capacity of old leaves remained constant throughout the study period, and there was no evidence that N was remobilized from Rubisco.     

Leaf physiology and sugar concentrations of transplanted Quercus rubra seedlings in relation to nutrient and water availability

Newly planted seedlings incur transplant stress resulting from poor root-soil contact, which limits access to soil moisture and nutrients and reduces growth for one or more growing seasons. Controlled release fertilizer (CRF) applied at planting may reduce transplant stress by augmenting rhizosphere nutrient availability yet with potential risk of root system damage due to elevated fertilizer salt concentrations, which may be further exacerbated by drought. Under controlled conditions, we examined northern red oak (Quercus rubra L.) leaf physiological parameters and soluble sugar concentrations in response to varying nutrient levels (via CRF application) and moisture availability gradients ranging from drought to flooding. Net photosynthetic rates, transpiration rates, and chlorophyll fluorescence parameters responded positively to CRF application, and no interactions were observed between CRF and moisture availability; however, CRF did not increase soluble sugar concentrations. No effects of short-term drought were observed, but flooding exerted a rapid negative influence on net photosynthetic rates, transpiration rates, and chlorophyll fluorescence parameters; flooding also elevated soluble sugar concentrations, indicative of disrupted carbon partitioning and a much greater sensitivity to root-zone hypoxia than to drought in this species. Lack of interactions between CRF application and soil moisture availability indicates relatively similar responses of fertilized seedlings across moisture gradients.     

Relationship between photosynthesis and leaf nitrogen concentration in ambient and elevated [CO2] in white birch seedlings

To study the effects of elevated CO2 concentration ([CO2]) on relationships between nitrogen (N) nutrition and foliar gas exchange parameters, white birch (Betula papyrifera Marsh.) seedlings were exposed to one of five N-supply regimes (10, 80, 150, 220, 290 mg N l(-1)) in either ambient [CO2] (360 micromol mol(-1)) or elevated [CO2] (720 micromol mol(-1)) in environment-controlled greenhouses. Foliar gas exchange and chlorophyll fluorescence were measured after 60 and 80 days of treatment. Photosynthesis showed a substantial down-regulation (up to 57%) in response to elevated [CO2] and the magnitude of the down-regulation generally decreased exponentially with increasing leaf N concentration. When measured at the growth [CO2], elevated [CO2] increased the overall rate of photosynthesis (P(n)) and instantaneous water-use efficiency (IWUE) by up to 69 and 236%, respectively, but decreased transpiration (E) and stomatal conductance (g(s)) in all N treatments. However, the degree of stimulation of photosynthesis by elevated [CO2] decreased as photosynthetic down-regulation increased from 60 days to 80 days of treatment. Elevated [CO2] significantly increased total photosynthetic electron transport in all N treatments at 60 days of treatment, but the effect was insignificant after 80 days of treatment. Both P(n) and IWUE generally increased with increasing leaf N concentration except at very high leaf N concentrations, where both P(n) and IWUE declined. The relationships of P(n) and IWUE with leaf N concentration were modeled with both a linear regression and a second-order polynomial function. Elevated [CO2] significantly and substantially increased the slope of the linear regression for IWUE, but had no significant effect on the slope for P(n). The optimal leaf N concentration for P(n) and IWUE derived from the polynomial function did not differ between the CO2 treatments when leaf N was expressed on a leaf area basis. However, the mass-based optimal leaf N concentration for P(n) was much lower in seedlings in elevated [CO2] than in ambient [CO2] (31.88 versus 37.00 mg g(-1)). Elevated [CO2] generally decreased mass-based leaf N concentration but had no significant effect on area-based leaf N concentration; however, maximum N concentration per unit leaf area was greater in elevated [CO2] than in ambient [CO2] (1.913 versus 1.547 g N m(-2)).     

Leaf photosynthesis, respiration and stomatal conductance in six Eucalyptus species native to mesic and xeric environments growing in a common garden

Trees adapted to mesic and xeric habits may differ in a suite of physiological responses that affect leaf-level carbon balance, including the relationship between photosynthesis (A) and respiration at night (R(n)). Understanding the factors that regulate physiological function in mesic and xeric species is critical for predicting changes in growth and distribution under changing climates. In this study, we examined the relationship between A and R(n), and leaf traits that may regulate A and R(n), in six Eucalyptus species native to mesic or xeric ecosystems, during two 24-h cycles in a common garden under high soil moisture. Peak A and R(n) generally were higher in xeric compared with mesic species. Across species, A and R(n) covaried, correlated with leaf mass per area, leaf N per unit area and daytime soluble sugar accumulation. A also covaried with g(s), which accounted for 93% of the variation in A within species. These results suggest that A and R(n) in these six Eucalyptus species were linked through leaf N and carbohydrates. Further, the relationship between A and R(n) across species suggests that differences in this relationship between mesic and xeric Eucalyptus species in their native habitats may be largely driven by environmental factors rather than inter-specific genetic variation.