Inhibition and acclimation of C(3) photosynthesis to moderate heat: a perspective from thermally contrasting genotypes of Acer rubrum (red maple)

Effects of moderate heat on growth and photosynthesis were investigated in two clonal genotypes of Acer rubrum L., originally collected from the thermally contrasting habitats of Florida and Minnesota, USA, and known in the horticultural trade for sensitivity and insensitivity to heat, respectively. Under both common garden and warm greenhouse conditions (day/night temperature of 33/25 degrees C), the Florida genotype exhibited more growth than the Minnesota genotype. To determine the physiological parameters associated with this response, plants were acclimated to ambient (27/25 degrees C) or moderately elevated (33/25 degrees C) temperatures for 21 days before measurement of net photosynthesis at temperatures ranging from 25 to 48 degrees C. In vivo measurements of gas exchange and chlorophyll a fluorescence of ambient-acclimated plants revealed that, compared with the Minnesota genotype, the Florida genotype maintained a higher photosynthetic rate, higher stomatal conductance, more open PSII reaction centers, a greater PSII quantum yield and a lower quantum requirement for photosystem II (phi(PSII)) per mole of CO(2) fixed (phi(CO(2) )) throughout the measurement temperature range. When both genotypes were acclimated at 33/25 degrees C and measured at 33 degrees C, analysis of the response of net photosynthesis to calculated intercellular CO(2) concentration indicated that the maximal rate of Rubisco carboxylation (V(cmax)) decreased more in the Minnesota genotype than in the Florida genotype in response to elevated temperature. Additionally, phi(PSII)/phi(CO(2) ) at 33 degrees C was markedly higher for Minnesota plants under photorespiratory conditions, but similar to Florida plants under non-photorespiratory conditions. The results indicate that the higher net photosynthetic rate at 33/25 degrees C of the Florida genotype compared with the Minnesota genotype could be a result of several mechanisms, including the maintenance of a higher V(cmax )and a more efficient quantum requirement of PSII per mole of CO(2) fixed, which is likely the result of lower photorespiration.     

Net CO(2) assimilation of cacao seedlings following dark chilling

Overnight exposure of cacao (Theobroma cacao L.) seedlings to chilling temperatures between 4.7 and 15.8 degrees C reduced net CO(2) assimilation rate (A) and stomatal conductance to water vapor (g(s)), with temperatures below 10 degrees C causing severe inhibition. Net CO(2) assimilation rates of chilled seedlings recovered to those of nonchilled plants within 7 days. No differences in daytime intercellular CO(2) concentration (c(i)) with overnight temperature were observed on the first day after the chilling treatment, which indicates that the reduction in photosynthesis was not caused by the reduction in stomatal conductance. However, c(i) of chilled plants was much less than that of nonchilled plants on the second day after treatment, which suggests that chilling caused a change in stomatal response to CO(2) concentration. Even 7 days after treatment, when A had recovered to control values, g(s) of chilled leaves was only approximately 70% that of controls. Chilling did not inhibit A through an effect on leaf water potential, which was higher in chilled plants than in unchilled plants.     

Acquired thermotolerance of jack pine, white spruce and black spruce seedlings

The acquired thermotolerance of first-year seedlings of jack pine (Pinus banksiana Lamb.) hardened at 36, 38, 40 or 42 degrees C for 90, 180 or 360 minutes and of black spruce (Picea mariana (Mill.) B.S.P.) hardened at 34, 36, 38 or 40 degrees C for 30, 90, 180 or 360 minutes was determined by comparison of needle damage to that of non-hardened seedlings (25 degrees C) following exposure to temperatures of 49 and 47.5 degrees C, respectively. Compared to seedlings kept at 25 degrees C, heat injury sustained from exposure to high temperatures was markedly reduced following hardening for 180 minutes at 36 and 38 degrees C in jack pine and black spruce, respectively. Increasing the exposure time at 36 degrees C in jack pine, and at 36 to 40 degrees C in black spruce, also reduced needle damage. The duration of increased thermotolerance was investigated in jack pine, black spruce and white spruce (Picea glauca (Moench) Voss) by comparing heat injury from high temperatures in non-hardened seedlings and in seedlings hardened at 38 degrees C for 180 minutes a day for either 1, 3 or 6 days. In all three species, the duration of acquired thermotolerance increased with the number of days of heat hardening. For jack pine and white spruce seedlings hardened at 38 degrees C for 6 days, increased thermotolerance persisted for at least 14 and 10 days, respectively, after the end of the hardening treatment. In contrast, the thermotolerance of black spruce seedlings hardened at 38 degrees C for 6 days remained elevated for only 4 days.     

Selection of white spruce families in the context of climate change: heat tolerance

To assess the responses and plasticity of white spruce seedlings (Picea glauca (Moench) Voss) to high temperatures, 12 open-pollinated families differing in growth performance were exposed to a 30-min heat treatment of 42, 44, 46, 48, or 50 degrees C with or without heat preconditioning at 38 degrees C for 5 h. Damage was evaluated based on chlorophyll fluorescence parameters after heat preconditioning, after the heat treatments and during a 7-day recovery period. Visible needle damage was also evaluated after the heat treatments and 14 days later. Chlorophyll fluorescence parameters indicated that seedlings subjected to a heat treatment of 42-43 degrees C lost the ability to phosphorylate and donate water to photosystem II (PSII). A heat treatment of 44-46 degrees C severely limited the ability of the seedlings to use NADPH and ATP in the Calvin cycle. Based on visible needle damage, families with superior height-growth performance were more sensitive to heat stress than families with intermediate or inferior height-growth performance. Moreover, families with superior height-growth performance had low photochemical efficiencies in the light (DeltaF/F(m)') after heat treatment. Heat preconditioning increased the thermotolerance of the seedlings. However, the data suggest that white spruce seedlings exhibiting fast-growing characteristics under present conditions may not grow as well at higher temperatures.     

黑河流域荒漠地区水分胁迫对梭梭苗木影响的试验研究(英文)

梭梭(HaloxylonAmmodendronBge,一种C4灌木)苗种植在15升的容器中,给予不同的水分胁迫处理,研究了其水分关系和气体交换特征。结果表明:当土壤水分含量大于11%时,梭梭苗有高的蒸腾量;土壤水分含量低于6%时,苗木就不能从土壤中吸取水分;很好供水的苗木的蒸腾量与潜在蒸发量成线型相关。气体交换测定发现,随着土壤水分含量的下降,造成了不同程度的气孔导度、叶蒸腾强度和光合作用的下降。对同一苗木而言,由于这个地区有高的水气压亏缺(VPD),很好和中度供水的苗木在气孔反应方面有较宽的范围,气孔在决定光合作用方面起着较小的作用,二者没有明显的线型相关关系。虽然水分胁迫使蒸腾速率比光合速率下降的更快,提高了水分利用效率,而较高的蒸发需求增加了蒸腾量,限制了光合作用,但是总的趋势是光合作用和蒸腾强度成线型相关。图6表2参15。     

Seasonal CO(2) assimilation and stomatal limitations in a Pinus taeda canopy

Net CO(2) assimilation (A(net)) of canopy leaves is the principal process governing carbon storage from the atmosphere in forests, but it has rarely been measured over multiple seasons and multiple years. I measured midday A(net) in the upper canopy of maturing loblolly pine (Pinus taeda L.) trees in the piedmont region of the southeastern USA on 146 sunny days over 36 months. Concurrent data for leaf conductance and photosynthetic CO(2) response curves (A(net)-C(i) curves) were used to estimate the relative importance of stomatal limitations to CO(2) assimilation in the field. In fully expanded current-year and 1-year-old needles, midday light-saturated A(net) was constant over much of the growing season (5-6 &mgr;mol CO(2) m(-2) s(-1)), except during drought periods. During the winter season (November-March), midday A(net) of overwintering needles varied in proportion to leaf temperature. Net CO(2) assimilation at light saturation occurred when daytime air temperatures exceeded 5-6 degrees C, as happened on more than 90% of the sunny winter days. In both age classes of foliage, winter carbon assimilation accounted for approximately 15% of the daily carbon assimilation on sunny days throughout the year, and was relatively insensitive to year-to-year differences in temperature during this season. However, strong stomatal limitations to A(net) occurred as a result of water stress associated with freezing cycles in winter. During the growing season, drought-induced water stress produced the largest year-to-year differences in seasonal CO(2) assimilation on sunny days. Seasonal A(net) was more drought sensitive in current-year needles than in 1-year-old needles. Relative stomatal limitations to daily integrated A(net) were approximately 40% over the growing season, and summer drought rather than high temperatures had the largest impact on summer A(net) and integrated annual CO(2) uptake in the upper crown. Despite significant stomatal limitations, a long duration of near-peak A(net) in the upper crown, particularly in 1-year-old needles, conferred high seasonal and annual carbon gain.     

Drought effects on hydraulic conductivity and xylem vulnerability to embolism in diverse species and provenances of Mediterranean cedars

We studied hydraulic traits of young plants of the Mediterranean cedar species Cedrus atlantica (Endl.) G. Manetti ex Carrière (Luberon, France), C. brevifolia (Hook. f.) Henry (Cyprus), C. libani A. Rich (Hadeth El Jebbe, Lebanon) and C. libani (Armut Alani, Turkey). With an optimum water supply, no major differences were observed among species or provenances in either stem hydraulic conductivity (Ks) or leaf specific conductivity (Kl) measured on the main shoot. A moderate soil drought applied for 10 weeks induced marked acclimation through a reduction in Ks, particularly in the Lebanese provenance of C. libani, and a decrease in tracheid lumen size in all species. Cedrus atlantica, which had the smallest tracheids, was the species most vulnerable to embolism: a 50% loss in hydraulic conductivity (PsiPLC50) occurred at a water potential of -4.4 MPa in the well-watered treatment, and at -6.0 MPa in the moderate drought treatment. In the other species, PsiPLC50 was unaffected by moderate soil drought, and only declined sharply at water potentials between -6.4 and -7.5 MPa in both irrigation treatments. During severe drought, Ks of twigs and stomatal conductance (g(s)) were measured simultaneously as leaf water potential declined. For all species, lower vulnerability to embolism based on loss of Ks was recorded on current-year twigs. The threshold for stomatal closure (10% of maximum g(s)) was reached at a predawn water potential (Psi(pd)) of -2.5 MPa in C. atlantica (Luberon) and at -3.1 MPa in C. libani (Lebanon), whereas the other provenance and species had intermediate Psi(pd) values. Cedrus brevifolia, with a Psi(pd) (-3.0 MPa) close to that of C. libani (Lebanon), had the highest stomatal conductance of the study species. The importance of a margin of safety between water potential causing stomatal closure and that causing xylem embolism induction is discussed.     

Thermal optima of photosynthetic functions and thermostability of photochemistry in cork oak seedlings

Temperature effects on photosynthesis were studied in seedlings of evergreen Mediterranean cork oak (Quercus suber L.). Responses to changes in temperature and the temperature optima of maximal carboxylation rate (V(cmax)) and maximal light-driven electron flux (J(max)) were estimated from gas exchange measurements and a leaf-level photosynthesis model. The estimated temperature optima were approximately 34 and 33 degrees C for V(cmax) and J(max), respectively, which fall within the lower range of temperature optima previously observed in deciduous tree species. The thermostability of the photosynthetic apparatus was estimated according to the temperature at which basal chlorophyll a fluorescence begins to increase (T(c)). The T(c) was highly variable, increasing from 42 to 51 degrees C when ambient temperature rose from 10 to 40 degrees C, and increasing from 44 to 54 degrees C with decreasing soil water availability while net CO(2) assimilation rate dropped to almost zero. When a heat shock was imposed, an additional small increase in T(c) was observed in drought-stressed and control seedlings. Maximal T(c) values following heat shock were about 56 degrees C, which, to our knowledge, are the highest values that have been observed in tree species. In conclusion, the intrinsic temperature responses of cork oak did not differ from those of other species (similar T(c) under ambient temperature and water availability, and relatively low thermal optima for photosynthetic capacity in seedlings grown at cool temperatures). However, the large ability of cork oak to acclimate to drought and elevated temperature may be an important factor in the tolerance of this evergreen Mediterranean species to summer drought and high temperatures.     

Nitrogen allocation and the fate of absorbed light in 21-year-old Pinus radiata

We investigated effects of nitrogen (N) fertilizer and canopy position on the allocation of N to Rubisco and chlorophyll as well as the distribution of absorbed light among thermal energy dissipation, photochemistry, net CO2 assimilation and alternative electron sinks such as the Mehler reaction and photorespiration. The relative reduction state of the primary quinone receptor of photosystem II (QA) was used as a surrogate for photosystem II (PSII) vulnerability to photoinactivation. Measurements were made on needles from the lower, mid and upper canopy of 21-year-old Pinus radiata D. Don trees grown with (N+) and without (N0) added N fertilizer. Rubisco was 45 to 60% higher in needles of N+ trees than in needles of N0 trees at all canopy positions. Chlorophyll was approximately 80% higher in lower- and mid-canopy needles of N+ trees than of N0 trees, but only approximately 20% higher in upper-canopy needles. Physiological differences between N+ and N0 trees were found only in the lower- and mid- canopy positions. Needles of N+ trees dissipated up to 30% less light energy as heat than needles of N0 trees and had correspondingly more reduced QA. Net CO2 assimilation and the proportions of electrons used by alternative electron sinks such as the Mehler reaction and photorespiration were unaffected by N treatment regardless of canopy position. We conclude that the application of N fertilizer mainly affected the biochemistry and light-use physiology in lower- and mid-canopy needles by increasing the amount of chlorophyll and hence the amount of light harvested. This, however, did not improve photochemistry or safe dissipation, but increased PSII vulnerability to photoinactivation, an effect with likely significant consequences during sunflecks or sudden gap formation.     

Contrasting effects of elevated carbon dioxide concentration and temperature on Rubisco activity,chlorophyll fluorescence,needle ultrastructure and secondary metabolites in conifer seedlings

Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) seedlings were grown for 50 days in growth chambers in an ambient or twice ambient carbon dioxide concentration ([CO2]) at a day/night temperature of 19/12 degrees C or 23/16 degrees C. Although elevated [CO2] (EC) had only slight effects on the growth parameters measured, elevated temperature (ET) increased above ground dry mass of both species. Among treatments, biomass accumulation of both species was greatest in the combined EC + ET treatment. The EC treatment induced thylakoid swelling and increased numbers of plastoglobuli observed in Scots pine needles. Although EC had little effect on Rubisco protein or N concentration of needles, ET had a large effect on N-containing compounds and enhanced N allocation from 1-year-old needles. Terpenoids were more responsive to EC and ET than total phenolics. Generally, terpene concentrations were reduced by EC and increased by ET. Increased terpenoid concentrations in response to ET might be associated with thermotolerance of photosynthesis. In Norway spruce, EC decreased total phenolic concentrations in needles, probably as a result of increased growth. We conclude that, in seedlings of these boreal species, the effects of elevated [CO2] on the studied parameters were small compared with the effects of elevated temperature.     

Photosynthetic traits around budbreak in pre-existing needles of Sakhalin spruce (Picea glehnii) seedlings grown under elevated CO2 concentration assessed by chlorophyll fluorescence measurements

To assess the effects of elevated CO(2) concentration ([CO(2)]) on the photosynthetic properties around spring budbreak, we monitored the total leaf sugar and starch content, and chlorophyll fluorescence in 1-year-old needles of Sakhalin spruce (Picea glehnii Masters) seedlings in relation to the timing of budbreak, grown in a phytotron under natural daylight at two [CO(2)] levels (ambient: 360?μmol mol(-1) and elevated: 720?μmol mol(-1)). Budbreak was accelerated by elevated [CO(2)] accompanied with earlier temporal declines in the quantum yield of PSII electron transport (Φ(PSII)) and photochemical quenching (q(L)). Plants grown under elevated [CO(2)] showed pre-budbreak leaf starch content twice as high with no significant difference in Φ(PSII) from ambient-CO(2)-grown plants when compared at the same measurement [CO(2)], i.e., 360 or 720?μmol mol(-1), suggesting that the enhanced pre-budbreak leaf starch accumulation might not cause down-regulation of photosynthesis in pre-existing needles under elevated [CO(2)]. Conversely, lower excitation pressure adjusted for the efficiency of PSII photochemistry ((1?-?q(P)) F(v)'/F(m)') was observed in plants grown under elevated [CO(2)] around budbreak when compared at their growth [CO(2)] (i.e., comparing (1?-?q(P)) F(v)'/F(m)' measured at 720?μmol mol(-1) in elevated-CO(2)-grown plants with that at 360?μmol mol(-1) in ambient-CO(2)-grown plants), which suggests lower rate of photoinactivation of PSII in the elevated-CO(2)-grown plants around spring budbreak. The degree of photoinhibition, as indicated by the overnight-dark-adapted F(v)/F(m), however, showed no difference between CO(2) treatments, thereby suggesting that photoprotection during the daytime or the repair of PSII at night was sufficient to alleviate differences in the rate of photoinactivation.     

Root water flow and growth of aspen (Populus tremuloides) at low root temperatures

Effects of root zone temperature on growth, shoot water relations, and root water flow were studied in 1-year-old aspen (Populus tremuloides Michx.) seedlings. Seedlings were grown in solution culture and exposed to day/night air temperatures of 22/16 degrees C and solution culture temperatures of 5, 10, or 20 degrees C for 28 days after bud flush. Compared with root growth at 20 degrees C, root growth was completely inhibited at 5 degrees C and inhibited by 97% at 10 degrees C. The 5 and 10 degrees C treatments severely reduced shoot growth, leaf size, and total leaf area. Root water flow was inhibited by the 5 and 10 degrees C treatments. However, when seedlings were grown for 28 days at 5 degrees C and root water flow was measured at 20 degrees C, there was an increase in flow rate. This increase in root water flow was similar in magnitude to the decrease in root water flow observed when seedlings were grown for 28 days at 20 degrees C and root water flow was measured at 5 degrees C. Reduced root water flow of seedlings grown at 5 and 10 degrees C resulted in decreased stomatal conductance, net assimilation, and shoot water potentials. Root water flow was positively correlated with leaf size, total leaf area, shoot length, and new root growth. Transferring seedlings from 5 to 20 degrees C for 24 h significantly increased root water flow, shoot water potential, and net photosynthesis, whereas transferring seedlings from 10 to 20 degrees C resulted in only a slightly increased shoot water potential. Transferring seedlings from 20 to 5 degrees C greatly reduced root water flow, stomatal conductance, and net photosynthesis, whereas shoot water potential decreased only slightly.     

Aluminum-induced decrease in CO2 assimilation in citrus seedlings is unaccompanied by decreased activities of key enzymes involved in CO2 assimilation

'Cleopatra' tangerine (Citrus reshni Hort. ex Tanaka) seedlings were irrigated daily for 8 weeks with 1/4 strength Hoagland's nutrient solution containing 0 (control) or 2 mM aluminum (Al). Leaves from Al-treated plants had decreased CO2 assimilation and stomatal conductance, but increased intercellular CO2 concentrations compared with control leaves. On a leaf area basis, 2 mM Al increased activities of key enzymes in the Calvin cycle, including ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco), NADP-glyceraldehyde-3-phosphate dehydrogenase (GAPDH), phosphoribulokinase (PRK), stromal fructose-1,6-bisphosphatase (FBPase), and a key enzyme in starch synthesis, ADP-glucose pyrophosphorylase (AGPase), compared with control leaves. Aluminum had no effect on cytosolic FBPase activity, but it decreased sucrose phosphate synthase (SPS) activity. Aluminum had no effect on area-based concentrations of carbohydrates, glucose-6-phosphate (G6P) and fructose 6-phosphate (F6P) or the G6P:F6P ratio, but it decreased the area-based concentration of 3-phosphoglycerate (PGA). Photochemical quenching coefficient (qP) and electron transport rate through PSII were greatly reduced by Al. Non-photochemical quenching coefficient (NPQ) was less affected by Al than qP and electron transport rate through PSII. We conclude that the reduced rate of CO2 assimilation in Al-treated leaves was probably caused by a combination of factors such as reduced electron transport rate through PSII, increased closure of PSII reaction centers and increased photorespiration.     

Seedlings of five boreal tree species differ in acclimation of net photosynthesis to elevated CO(2) and temperature

Biochemical models of photosynthesis suggest that rising temperatures will increase rates of net carbon dioxide assimilation and enhance plant responses to increasing atmospheric concentrations of CO(2). We tested this hypothesis by evaluating acclimation and ontogenetic drift in net photosynthesis in seedlings of five boreal tree species grown at 370 and 580 &mgr;mol mol(-1) CO(2) in combination with day/night temperatures of 18/12, 21/15, 24/18, 27/21, and 30/24 degrees C. Leaf-area-based rates of net photosynthesis increased between 13 and 36% among species in plants grown and measured in elevated CO(2) compared to ambient CO(2). These CO(2)-induced increases in net photosynthesis were greater for slower-growing Picea mariana (Mill.) B.S.P., Pinus banksiana Lamb., and Larix laricina (Du Roi) K. Koch than for faster-growing Populus tremuloides Michx. and Betula papyrifera Marsh., paralleling longer-term growth differences between CO(2) treatments. Measures at common CO(2) concentrations revealed that net photosynthesis was down-regulated in plants grown at elevated CO(2). In situ leaf gas exchange rates varied minimally across temperature treatments and, contrary to predictions, increasing growth temperatures did not enhance the response of net photosynthesis to elevated CO(2) in four of the five species. Overall, the species exhibited declines in specific leaf area and leaf nitrogen concentration, and increases in total nonstructural carbohydrates in response to CO(2) enrichment. Consequently, the elevated CO(2) treatment enhanced rates of net photosynthesis much more when expressed on a leaf area basis (25%) than when expressed on a leaf mass basis (10%). In all species, rates of leaf net CO(2) exchange exhibited modest declines with increasing plant size through ontogeny. Among the conifers, enhancements of photosynthetic rates in elevated CO(2) were sustained through time across a wide range of plant sizes. In contrast, for Populus tremuloides and B. papyrifera, mass-based photosynthetic rates did not differ between CO(2) treatments. Overall, net photosynthetic rates were highly correlated with relative growth rate as it varied among species and treatment combinations through time. We conclude that interspecific variation may be a more important determinant of photosynthetic response to CO(2) than temperature.     

Photosynthetic decline and pigment loss during autumn foliar senescence in western larch (Larix occidentalis)

We measured needle pigment content and photosynthetic rates of 1-year-old western larch (Larix occidentalis Nutt.) during autumn foliar senescence. Chlorophyll (Chl) and carotenoid (xanthophyll + b-carotene) contents of needles declined 11 and 17%, respectively, before CO(2) assimilation rate began to decline. Chlorophyll a/b ratio, Chl/carotenoid ratio, photochemical efficiency (F(v)/F(m)), and photochemical quenching did not begin to decline until late in senescence. Internal CO(2)/ambient CO(2) did not change during needle yellowing. In seedlings in warmed soil (average 3 degrees C above natural conditions), the decline in needle chlorophyll content was delayed by 10 days and the decline in CO(2) assimilation rate was delayed by 5 days, compared with seedlings in soil at ambient temperature. In seedlings exposed to an extended 16-h photoperiod, the decline in needle chlorophyll content was delayed by 32 days, and the decline in CO(2) assimilation rate was delayed by 21 days, compared with seedlings exposed to natural day lengths. In addition to delaying the onset of needle senescence, the treatments affected the sequence of events during senescence. Differences among treatment groups provide evidence that the onset of pigment loss and photosynthetic decline and the sequence of events during needle senescence are affected by soil temperature and day length.     

Daily time course of whole-shoot gas exchange rates in two drought-exposed Mediterranean shrubs

Effects of drought on water relations, whole-shoot gas-exchange characteristics, and pigment and zeatin concentrations were investigated in the Mediterranean shrubs rosemary (Rosmarinus officinalis L.) and lavender (Lavandula stoechas L.). Two-year-old, greenhouse-grown plants were placed in a whole-shoot gas-exchange measurement system and subjected to 10 days of drought, resulting in severe water stress, and then re-watered for 5 days in order to study their recovery. Water stress resulted in a significant decline in maximum whole-shoot net CO2 assimilation rates (An) for both species that was associated with reductions in leaf area and stomatal conductance. Because shoot dark respiration rate (Rd) was less sensitive to water stress than An, shoot Rd/An ratio increased from about 15 to 95% during water stress. No major changes in chlorophyll and carotenoid concentrations of rosemary leaves were observed during the experiments, but chlorophyll and carotenoid concentrations fell significantly in water-stressed lavender leaves. Zeatin concentrations were higher in rosemary leaves than in lavender leaves during water stress. After re-watering, whole-shoot An and Rd rapidly recovered to their pre-drought rates.     

Higher growth temperatures decreased net carbon assimilation and biomass accumulation of northern red oak seedlings near the southern limit of the species range

If an increase in temperature will limit the growth of a species, it will be in the warmest portion of the species distribution. Therefore, in this study we examined the effects of elevated temperature on net carbon assimilation and biomass production of northern red oak (Quercus rubra L.) seedlings grown near the southern limit of the species distribution. Seedlings were grown in chambers in elevated CO(2) (700 μmol mol(-1)) at three temperature conditions, ambient (tracking diurnal and seasonal variation in outdoor temperature), ambient +3 °C and ambient +6 °C, which produced mean growing season temperatures of 23, 26 and 29 °C, respectively. A group of seedlings was also grown in ambient [CO(2)] and ambient temperature as a check of the growth response to elevated [CO(2)]. Net photosynthesis and leaf respiration, photosynthetic capacity (V(cmax), J(max) and triose phosphate utilization (TPU)) and chlorophyll fluorescence, as well as seedling height, diameter and biomass, were measured during one growing season. Higher growth temperatures reduced net photosynthesis, increased respiration and reduced height, diameter and biomass production. Maximum net photosynthesis at saturating [CO(2)] and maximum rate of electron transport (J(max)) were lowest throughout the growing season in seedlings grown in the highest temperature regime. These parameters were also lower in June, but not in July or September, in seedlings grown at +3 °C above ambient, compared with those grown in ambient temperature, indicating no impairment of photosynthetic capacity with a moderate increase in air temperature. An unusual and potentially important observation was that foliar respiration did not acclimate to growth temperature, resulting in substantially higher leaf respiration at the higher growth temperatures. Lower net carbon assimilation was correlated with lower growth at higher temperatures. Total biomass at the end of the growing season decreased in direct proportion to the increase in growth temperature, declining by 6% per 1 °C increase in mean growing season temperature. Our observations suggest that increases in air temperature above current ambient conditions will be detrimental to Q. rubra seedlings growing near the southern limit of the species range.     

Effects of soil drought stress on photosynthetic gas exchange traits and chlorophyll fluorescence in Forsythia suspensa

To clarify the changes in plant photosynthesis and mechanisms underlying those responses to gradually increasing soil drought stress and reveal quantitative relationships between photosynthesis and soil moisture,soil water conditions were controlled in greenhouse pot experiments using 2-year-old seedlings of Forsythia suspensa(Thunb.) Vahl. Photosynthetic gas exchange and chlorophyll fluorescence variables were measured and analyzed under 13 gradients of soil water content. Net photosynthetic rate(PN), stomatal conductance(gs), and water-use efficiency(WUE) in the seedlings exhibited a clear threshold response to the relative soil water content(RSWC). The highest PNand WUEoccurred at RSWCof51.84 and 64.10%, respectively. Both PNand WUEwere higher than the average levels at 39.79% B RSWCB 73.04%. When RSWCdecreased from 51.84 to 37.52%,PN, gs, and the intercellular CO2 concentration(Ci)markedly decreased with increasing drought stress; the corresponding stomatal limitation(Ls) substantially increased, and nonphotochemical quenching(NPQ) also tended to increase, indicating that within this range of soil water content, excessive excitation energy was dispersed from photosystem II(PSII) in the form of heat, and the reduction in PNwas primarily due to stomatal limitation.While RSWCdecreased below 37.52%, there were significant decreases in the maximal quantum yield of PSII photochemistry(Fv/Fm) and the effective quantum yield of PSII photochemistry(UPSII), photochemical quenching(qP), and NPQ; in contrast, minimal fluorescence yield of the dark-adapted state(F0) increased markedly. Thus,the major limiting factor for the PNreduction changed to a nonstomatal limitation due to PSII damage. Therefore, an RSWCof 37.52% is the maximum allowable water deficit for the normal growth of seedlings of F. suspensa, and a water content lower than this level should be avoided in field soil water management. Water contents should be maintained in the range of 39.79% B RSWCB 73.04% to ensure normal function of the photosynthetic apparatus and high levels of photosynthesis and efficiency in F.suspensa.     

Excitation energy partitioning and quenching during cold acclimation in Scots pine

We studied the influence of two irradiances on cold acclimation and recovery of photosynthesis in Scots pine (Pinus sylvestris L.) seedlings to assess mechanisms for quenching the excess energy captured by the photosynthetic apparatus. A shift in temperature from 20 to 5 degrees C caused a greater decrease in photosynthetic activity, measured by chlorophyll fluorescence and oxygen evolution, in plants exposed to moderate light (350 micromol m(-2) s(-1)) than in shaded plants (50 micromol m(-2) s(-1)). In response to the temperature shift, maximal photochemical efficiency of photosystem II (PSII), measured as the ratio of variable to maximal chlorophyll fluorescence (Fv/Fm) of dark-adapted samples, decreased to 70% in exposed seedlings, whereas shaded seedlings maintained Fv/Fm close to initial values. After a further temperature decrease to -5 degrees C, only 8% of initial Fv/Fm remained in exposed plants, whereas shaded plants retained 40% of initial Fv/Fm. Seven days after transfer from -5 to 20 degrees C, recovery of photochemical efficiency was more complete in the shaded plants than in the exposed plants (87 and 65% of the initial Fv/Fm value, respectively). In response to cold stress, the estimated functional absorption cross section per remaining PSII reaction center increased at both irradiances, but the increase was more pronounced in exposed seedlings. Estimates of energy partitioning in the needles showed a much higher dissipative component in the exposed seedlings at low temperatures, pointing to stronger development of non-photochemical quenching at moderate irradiances. The de-epoxidation state of the xanthophyll cycle pigments increased in exposed seedlings at 5 degrees C, contributing to the quenching capacity, whereas significant de-epoxidation in the shaded plants was observed only when temperatures decreased to -5 degrees C. Thermoluminescence (TL) measurements of PSII revealed that charge recombinations between the second oxidation state of Mn-cluster S2 and the semireduced secondary electron acceptor quinone Q(B)- (S2Q(B)-) were shifted to lower temperatures in cold-acclimated seedlings compared with control seedlings and this effect depended on irradiance. Concomitant with this, cold-acclimated seedlings demonstrated a significant shift in the S2 recombination with primary acceptor Q(A)- (S2Q(A)-) characteristic TL emission peak to higher temperatures, thus narrowing the redox potential gap between S2Q(B)- and S2Q(A)-, which might result in increased probability for non-radiative radical pair recombination between the PSII reaction center chlorophyll a (P680+) and Q(A)- (P680+)Q(A)-) (reaction center quenching) in cold-acclimated seedlings. In Scots pine seedlings, mechanisms of quenching excess light energy in winter therefore involve light-dependent regulation of reaction center content and both reaction center-based and antenna-based quenching of excess light energy, enabling them to withstand high excitation pressure under northern winter conditions.     

Partial shoot removal increases net CO(2) assimilation and alters water relations of Citrus seedlings

Effects of defoliation on partial shoot removal by decapitation on seedling growth, water use and net gas exchange of remaining basal leaves, were examined in Citrus spp. Shoot and root growth rates were manipulated to test for effects of growth demands on net gas exchange. Partially defoliated plants had higher leaf pressure potentials, root conductivities and rates of water use than intact control plants. Shoot regrowth occurred at the expense of root loss. Basal leaves on defoliated plants consistently had higher rates of CO(2) assimilation (A) than leaves on intact plants. Stomatal conductance (g(s)) changed little after defoliation so the higher A of leaves on defoliated plants lowered the ratio of intercellular to ambient CO(2) concentration (C(i)/C(a)) in the mesophyll. In some cases, g(s) increased with A in defoliated plants but C(i)/C(a) was not affected. Stomatal conductance only limited A when intact seedlings were stressed by root confinement in small pots or when leaves were exposed to high vapor pressure deficits during gas exchange measurements. Increased carbon demand for shoot regrowth increased photosynthetic capacity and was more important than stomatal responses in determining A after partial shoot loss.