Photosynthetic response of white spruce families to drought stress

In the context of climate change, an increased frequency of drought stresses might occur at a regional scale in boreal forests. To assess photosynthetic responses to drought treatment, seedlings of 12 open-pollinated families of white spruce (Picea glauca (Moench) Voss) differing in their growth performance were grown in a controlled environment. Gas exchange and chlorophyll fluorescence parameters as well as shoot xylem water potential (WP) were measured for 21 successive days after watering was stopped. Net photosynthesis decreased as stomatal conductance decreased. Net photosynthesis was not affected by drought until WP reached –2.0 MPa when stomata were closed. Initial fluorescence (F and basic fluorescence after induction (F00) were not affected by drought. A progressive decrease in maximal (Fm) and variable fluorescences (Fv), maximum photosystem II (PS II) efficiency (Fv = Fm), effective quantum yield of PS II (FII), photochemical efficiency of open PS II (Fp), and photochemical quenching (qP) was observed at WP < - 1.0 MPa, whereas non-photochemical quenching (qN) remained high throughout the drought treatment. White spruce families with inferior growth performance showed higher values of Fm, Fv, Fv = Fm, Fp, and qN at WP< - 2.0MPa. The results indicated that chlorophyll fluorescence variables can be used as drought markers in relation to present or predicted climate conditions. These could be used for selecting planting stock adapted to drought periods or dry environments. These markers showed that slow-growing genotypes are better adapted to drought conditions than intermediate or fast-growing genotypes in present and predicted drought conditions.     

Effects of cone-induction treatments on black spruce (Picea mariana) current-year needle development and gas exchange properties

Both drought and root pruning (RP) increased the number of cones induced when black spruce (Picea mariana (Mill.) B.S.P.) grafts were injected with gibberellins A(4/7) (GA), but their effects on predawn shoot water potential and current-year needle development differed. Drought decreased predawn shoot water potential (Psi(pd)), but only during the period when irrigation was withheld, and it had no effect on the growth or gas exchange properties of current-year needles. Conversely, root pruning had little effect on Psi(pd), but it resulted in trees with smaller current-year needles that had lower nitrogen and chlorophyll concentrations and reduced rates of gas exchange up to the later stages of shoot elongation compared with needles of control trees. These findings are discussed in relation to potential effects on the development of induced cones in the following growth cycle.     

Effects of drought preconditioning on thermotolerance of photosystem II and susceptibility of photosynthesis to heat stress in cedar seedlings

Changes in photosystem II (PSII) thermotolerance during drought and recovery were studied under controlled conditions in three Mediterranean cedar species (Cedrus brevifolia Henry, C. libani Loudon and C. atlantica Manetti). The temperature at which the quantum yield of PSII photochemistry was reduced by 15% of its value at 25 degrees C was 3 to 4 degrees C higher in drought-treated plants than in well-watered plants. The drought-induced increase in PSII thermotolerance was already evident 8 days after water had been withheld from the seedlings, when net CO(2) assimilation was still at 80% of its initial value, and was visible for up to 12 days after re-watering. When seedlings of the three species were exposed to temperatures above 45 degrees C for 5 h, both maximal quantum yield of PSII photochemistry and net CO(2) assimilation rate were significantly reduced in unconditioned seedlings, whereas drought-preconditioned seedlings were almost unaffected by the heat treatment. Drought-preconditioned seedlings still exhibited a higher tolerance to heat stress than unconditioned seedlings 60 days after re-watering, although the transient, drought-induced osmotic adjustment had fully disappeared. Among species, C. atlantica was the most heat sensitive, whereas the heat treatment had no significant effect on the parameters measured in C. brevifolia.     

Physiological response to drought in radiata pine: phytohormone implication at leaf level

Pinus radiata D. Don is one of the most abundant species in the north of Spain. Knowledge of drought response mechanisms is essential to guarantee plantation survival under reduced water supply as predicted in the future. Tolerance mechanisms are being studied in breeding programs, because information on such mechanisms can be used for genotype selection. In this paper, we analyze the changes of leaf water potential, hydraulic conductance (K(leaf)), stomatal conductance and phytohormones under drought in P. radiata breeds (O1, O2, O3, O4, O5 and O6) from different climatology areas, hypothesizing that they could show variable drought tolerance. As a primary signal, drought decreased cytokinin (zeatin and zeatin riboside-Z?+?ZR) levels in needles parallel to K(leaf) and gas exchange. When Z?+?ZR decreased by 65%, indole-3-acetic acid (IAA) and abscisic acid (ABA) accumulation started as a second signal and increments were higher for IAA than for ABA. When plants decreased by 80%, Z?+?ZR and K(leaf) doubled their ABA and IAA levels, the photosystem II yield decreased and the electrolyte leakage increased. At the end of the drought period, less tolerant breeds increased IAA over 10-fold compared with controls. External damage also induced jasmonic acid accumulation in all breeds except in O5 (P. radiata var. radiata?×?var. cedrosensis), which accumulated salicylic acid as a defense mechanism. After rewatering, only the most tolerant plants recovered their K(leaf,) perhaps due to an IAA decrease and 1-aminocyclopropane-1-carboxylic acid maintenance. From all phytohormones, IAA was the most representative 'water deficit signal' in P. radiata.     

Paclobutrazol affects the resistance of black spruce to high light and thermal stress

Detached needles from 20-week-old black spruce (Picea mariana (Mill.) B.S.P.) seedlings root-drenched with 60 mg of paclobutrazol were exposed to two temperatures (22 and 50 degrees C) and two light treatments (100 and 1900 &mgr;mol m(-2) s(-1) PAR) in a factorial combination for 4 h in vitro. Mean dry weights of individual needles from paclobutrazol-treated plants were approximately 1.9 times heavier than that of needles from untreated controls at 22 degrees C, but no differences were observed following incubation at 50 degrees C. Numbers of cells per needle remained constant in all treatments. Chlorophyll and carotenoid contents per needle were higher in seedlings treated with paclobutrazol than in untreated control seedlings, and the differences were most pronounced in the high temperature plus high light treatment. In low light at 50 degrees C, quantum efficiency of photosystem II was 45% higher in needles of paclobutrazol-treated seedlings than in needles of untreated control seedlings, but quantum efficiency of needles from treated seedlings declined when needles were exposed to high light at either temperature. Peroxidase and superoxide dismutase activities were up-regulated by paclobutrazol, whereas catalase activities were depressed and no significant differences were observed between treated and control needles at 50 degrees C in either light treatment. Paclobutrazol treatment did not moderate the depressive effects of high temperature on total soluble protein or on the activity of ribulose-1,5-bisphosphate carboxylase. In contrast, high activities of phosphoenolpyruvate carboxylase were maintained in paclobutrazol-treated needles under all stress conditions, whereas large losses in activity were recorded in untreated needles at 50 degrees C. Collectively, these observations suggest that paclobutrazol treatment may convey resistance to excessive light and high temperatures by increasing the potential of conifers to limit damage caused by oxidative stress.     

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.     

The response of Pinus sylvestris to drought: stomatal control of transpiration and hydraulic conductance

We investigated the impact of drought on the physiology of 41-year-old Scots pine (Pinus sylvestris L.) in central Scotland. Measurements were made of the seasonal course of transpiration, canopy stomatal conductance, needle water potential, xylem water content, soil-to-needle hydraulic resistance, and growth. Comparison was made between drought-treated plots and those receiving average precipitation. In response to drought, transpiration rate declined once volumetric water content (VWC) over the top 20 cm of soil reached a threshold value of 12%. Thereafter, transpiration was a near linear function of soil water content. As the soil water deficit developed, the hydraulic resistance between soil and needles increased by a factor of three as predawn needle water potential declined from -0.54 to -0.71 MPa. A small but significant increase in xylem embolism was detected in 1-year-old shoots. Stomatal control of transpiration prevented needle water potential from declining below -1.5 MPa. Basal area, and shoot and needle growth were significantly reduced in the drought treatment. In the year following the drought, canopy stomatal conductance and soil-to-needle hydraulic resistance recovered. Current-year needle extension recovered, but a significant reduction in basal area increment was evident one year after the drought. The results suggest that, in response to soil water deficit, mature Scots pine closes its stomata sufficiently to prevent the development of substantial xylem embolism. Reduced growth in the year after a severe soil water deficit is most likely to be the result of reduced assimilation in the year of the drought, rather than to any residual embolism carried over from one year to the next.     

Needle growth in Sitka spruce (Picea sitchensis): effects of nutrient deficiency and needle position within shoots

Needle development and shoot growth were studied in 14- and 20-year-old Sitka spruce (Picea sitchensis (Bong.) Carr.) planted on an oligotrophic peat and fertilized with N, P and K (control), N and P (-K), N and K (-P) or unfertilized (-NPK). Shoot extension, needle number, and the size and number of needle cells were observed throughout the season. Beginning with bud burst in early May, needle growth continued for nine weeks in the control treatment. Needle growth in the mineral deficiency treatments terminated at the same time as in the controls, although it did not begin until up to three weeks later than in the control trees. Needles developed acropetally along the shoot, with basal needles completing their development first. Cell division in needles of control trees lasted about 3 weeks, with cell expansion continuing for a further 3 weeks. Initial mean cell cycle times were shortest for proximal needles following bud burst, but lengthened as the season progressed. Cell number increased from 6600 in the primordia to 200 000 in mature needles. Final needle dimensions and cell number varied according to the position of the needle on the shoot. The largest needles with the most cells were near the middle of the shoot. Relative to the controls, all mineral deficiency treatments reduced shoot length, bud and needle dimensions, and needle cell number, particularly the -P treatment. Potassium and P deficiency treatments reduced primordial cell number by up to 42%, whereas final mean cell size was 30% greater in the -P treatment, 17% greater in the -NPK treatment, but 14% smaller in the -K treatment. Mean cell size was constant for needles at all positions in any particular treatment at any time, so that final needle size was determined by cell number alone. Needle dry weight/fresh weight ratio continued to increase until early August, with significantly higher ratios in the -K and -P treatments than in the controls.     

Drought-induced oxidative stress in Canarian laurel forest tree species growing under controlled conditions

We studied photoprotection and antioxidative protection in the three major species of the Canarian laurel forest (Laurus azorica (Seub.) Franco, Persea indica (L.) K. Spreng and Myrica faya Aiton). Trees were exposed to drought under controlled conditions by withholding water until leaf relative water content (RWC) reached 50-55%. Drought reduced photosynthetic rate (P(N)) and was associated with decreased quantum yield of photosystem II (PSII) electron transport and increased non-photochemical quenching (NPQ) in L. azorica and M. faya, but did not increase NPQ in P. indica. Drought-treated trees of L. azorica had the highest de-epoxidation state (DPS) of the xanthophyll cycle and the highest zeaxanthin (Z) concentration, suggesting that this species had more effective photoprotective mechanisms than M. faya and P. indica. Moreover, beta-carotene remained unaltered in L. azorica trees during drought, suggesting that the chloroplasts of this species are better protected against oxidative stress than those of M. faya and P. indica. Increased antioxidation by ascorbate peroxidase, superoxide dismutase and glutathione reductase in L. azorica removed activated oxygen species (AOS) generated during drought treatment. Although M. faya was able to increase its energy dissipation rate by forming Z and thus increasing the DPS of the xanthophyll cycle, it did not respond to drought-induced oxidative stress with the result that beta-carotene degradation occurred. Persea indica did not activate an energy dissipation mechanism in response to drought treatment, hence formation of AOS was likely high in the drought-treated trees. In general, L. azorica was most resistant and P. indica most sensitive to photoinhibition and oxidative stress during drought.     

Stomatal limitation to CO2 assimilation and down-regulation of photosynthesis in Quercus ilex resprouts in response to slowly imposed drought

Holm oak (Quercus ilex L.) is native to hot, dry Mediterranean forests where limited water availability often reduces photosynthesis in many species, and forest fires are frequent. Holm oaks resprout after a disturbance, with improved photosynthetic activity and water relations compared with unburned plants. To better understand the role of water availability in this improvement, watering was withheld from container-grown plants, either intact (controls) or resprouts after excision of the shoot, to gradually obtain a wide range of soil water availabilities. At high water availability, gas exchange rates did not differ between controls and resprouts. At moderate soil dryness, net photosynthesis of control plants decreased as a result of increased stomatal limitation, whereas gas exchange rates of resprouts, which had higher midday and predawn leaf water potentials, were unchanged. Under severe drought, resprouts showed a less marked decline in gas exchange than controls and maintained photosystem II integrity, as indicated by chlorophyll fluorescence measurements. Photosynthesis was down-regulated in both plant types in response to reduced CO2 availability caused by high stomatal limitation. Lower non-stomatal limitations in resprouts than in control plants, as evidenced by higher carboxylation velocity and the capacity for ribulose-1,5-bisphosphate regeneration, conferred greater drought resistance under external constraints similar to summer conditions at midday.     

土壤温度和水分变化对川西云杉幼苗氮和磷含量的影响

【目的】研究不同梯度的土壤温度和水分对川西云杉幼苗生长性状和各器官氮和磷含量的影响,以期为全球气候变暖背景下解释川西云杉树线形成的原因提供参考和数据积累。【方法】以5年生川西云杉幼苗为试验材料,采用人工气候室结合嵌套设计,设置5个土壤温度梯度(2、7、12、17、22℃)和3个土壤水分梯度(干旱处理、正常水分含量处理、饱和水分含量处理)。每处理9株幼苗,共135株幼苗。实验处理4个月后,测定并比较分析不同梯度的土壤温度和水分对幼苗的生长性状、各器官干物质含量、各器官全氮、全磷浓度和含量以及土壤全氮和全磷浓度的影响。【结果】土壤温度处理对幼苗基径和株高生长量均无显著影响,而土壤水分处理对幼苗株高生长量有显著影响;在2℃和7℃土壤温度干旱处理下显著降低了幼苗的株高生长量,但随着土壤温度的升高其影响效应不显著。土壤温度处理对土壤氮和磷浓度无显著影响,而干旱处理显著升高了土壤氮和磷浓度。川西云杉幼苗各器官的氮和磷浓度以及当年生叶氮含量随土壤温度降低显著降低;干旱和饱和水分处理显著降低了当年生叶和当年生枝的氮浓度,饱和水分处理显著降低了当年生叶的磷浓度,干旱处理显著降低了当年生枝、茎和根的磷浓度,干旱和饱和水分处理显著降低了当年生叶和根的氮和磷含量,且随着土壤温度升高影响效应更显著。【结论】在短期内,土壤低温对川西云杉幼苗的生长性状没有明显的制约作用,但对川西云杉幼苗各器官的氮和磷浓度及含量影响显著,尤其是当年生叶和根的氮和磷浓度及含量。在川西地区,低温、干旱等极端气候胁迫导致的云杉幼苗氮、磷含量的不足很可能是限制川西云杉垂直分布的重要因素。此外,土壤温度和水分处理存在显著的交互作用,随着土壤温度的降低,水分胁迫对幼苗各器官氮和磷含量的影响由显著变得不再显著,说明随着海拔升高,与水分因子相比,土壤低温成为造成云杉各器官营养元素亏缺的主导因子。     

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.     

Reduction in turgid water volume in jack pine,white spruce and black spruce in response to drought and paclobutrazol

Significant reductions in needle water content were observed in white spruce (Picea glauca (Moench) Voss), black spruce (Picea mariana (Mill) B.S.P.), and jack pine (Pinus banksiana Lamb.) seedlings in response to a 10-day drought, although turgor was apparently maintained. When the seedlings were re-watered after the drought, jack pine needles regained their original saturated volume, whereas white spruce and black spruce needles did not. Significant drought-induced reductions in turgor-loss volume (i.e., tissue volume at the point of turgor loss) were observed in shoots of all three species, especially jack pine. Repeated exposure to 7 days of drought or treatment with the cytochrome P(450) inhibitor, paclobutrazol ((2RS,3RS)-1-(4-chlorophenyl)-4,4-dimethyl-2-(1,2,4-triazol-1-yl)-pentan-3-ol), reduced seedling height relative to that of untreated controls in all three species. The reductions in saturated and turgor-loss needle volumes in the paclobutrazol-treated seedlings were comparable with those of seedlings subjected to a 10-day drought. The treatment-induced reductions in shoot and needle water contents enabled seedlings to maintain turgor with tissue volumes close to, or below, the turgor-loss volume of untreated seedlings. Paclobutrazol-treated seedlings subsequently survived drought treatments that were lethal to untreated seedlings.     

Acclimation of shade-developed leaves on saplings exposed to late-season canopy gaps

We hypothesized that photoinhibition of shade-developed leaves of deciduous hardwood saplings would limit their ability to acclimate photosynthetically to increased irradiance, and we predicted that shade-tolerant sugar maple (Acer saccharum Marsh.) would be more susceptible to photoinhibition than intermediately shade-tolerant red oak (Quercus rubra L.). After four weeks in a canopy gap, photosynthetic rates of shade-developed leaves of both species had increased in response to the increase in irradiance, although final acclimation was more complete in red oak. However, photoinhibition occurred in both species, as indicated by short-term reductions in maximum rates of net photosynthesis and the quantum yield of oxygen evolution, and longer-term reductions in the efficiency of excitation energy capture by open photosystem II (PSII) reaction centers (dark-adapted F(v)/F(m)) and the quantum yield of PSII in the light (phi(PSII)). The magnitude and duration of this decrease were greater in sugar maple than in red oak, suggesting greater susceptibility to photoinhibition in sugar maple. Photoinhibition may have resulted from photodamage, but it may also have involved sustained rates of photoprotective energy dissipation (especially in red oak). Photosynthetic acclimation also appeared to be linked to an ability to increase leaf nitrogen content. Limited photosynthetic acclimation in shade-developed sugar maple leaves may reflect a trade-off between shade-tolerance and rapid acclimation to a canopy gap.     

Responses of Picea mariana to elevated CO2 concentration during growth, cold hardening and dehardening: phenology, cold tolerance, photosynthesis and growth

Seedlings from a northern and a southern provenance of black spruce (Picea mariana Mill. BSP) from eastern Canada were exposed to 37 or 71 Pa of carbon dioxide (CO2) during growth, cold hardening and dehardening in a greenhouse. Bud phenology, cold tolerance and photosynthetic efficiency were assessed during the growing and over-wintering periods. Bud set occurred earlier in elevated [CO2] than in ambient [CO2], but it was later in the southern provenance than in the northern provenance. An increase in seedling cold tolerance in early fall was related to early bud set in elevated [CO2]. Maximal photosystem II (PSII) photochemical efficiency (F(v)/F(m)), effective quantum yield (phi(PSII)), photochemical quenching (q(P)), light-saturated photosynthesis (Amax), apparent quantum efficiency (alpha'), light-saturated rate of carboxylation (Vcmax) and electron transport (Jmax) decreased during hardening and recovered during dehardening. Although Amax and alpha' were higher in elevated [CO2] when measured at the growth [CO2], down-regulation of photosynthesis occurred in elevated [CO2] as shown by lower F(v)/F(m), phi(PSII), Vcmax and Jmax. Elevated [CO2] reduced gene expression of the small subunit of Rubisco and also decreased chlorophyll a/chlorophyll b ratio and nitrogen concentration in needles, confirming our observation of down-regulation of photosynthesis. Elevated [CO2] increased the CO2 diffusion gradient and decreased photorespiration, which may have contributed to enhance Amax despite down-regulation of photosynthesis. Total seedling dry mass was higher in elevated [CO2] than in ambient [CO2] at the end of the growing season. However, because of earlier bud formation and cold hardening, and down-regulation of photosynthesis during fall and winter in elevated [CO2], the treatment difference in dry mass increment was less by the end of the winter than during the growing season. Differences in photosynthetic rate observed during fall, winter and spring account for the inter-annual variations in carbon assimilation of black spruce seedlings: our results demonstrate that these variations need to be considered in carbon budget studies.     

美国山核桃无性系叶绿素的荧光特性

研究了12个不同品种美国山核桃无性系嫁接苗的叶绿素荧光特性.结果表明:不同品种间的初始荧光(Fo)、最大荧光(Fm)、PS II原初光能转化效率(Fv/Fm)、PS II潜在活性(Fv/Fo)、非光化学猝灭系数(qN)均存在显著或极显著差异;光合量子产额(Y ield)和光化学猝灭系数(qP)未达显著差异.初步聚类分析表明,不同无性系嫁接苗中的所有西部品种和国内优系金华1号具有较好的光合性能.     

The effects of ultraviolet-B radiation on photosynthesis of different aged needles in field-grown loblolly pine

We examined the effect of supplemental UV-B radiation (290-320 nm) on photosynthetic characteristics of different aged needles of 3-year-old, field-grown loblolly pine (Pinus taeda L.). Needles in four age classes were examined: I, most recently fully expanded, year 3; II, first flush, year 3; III, final flush, year 2; and IV, oldest needles still present, year 2. Enhanced UV-B radiation caused a statistically significant decrease (6%) in the ratio of variable to maximum fluorescence (F(v)/F(m)) following dark adaptation only in needles from the youngest age class, suggesting transient damage to photosynthesis. However, no effects of enhanced UV-B radiation on other instantaneous measures of photosynthesis, including maximum photosynthesis, apparent quantum yield and dark respiration, were seen for needles of any age. Foliar nitrogen concentration was unaffected by UV-B treatment. However, the (13)C/(12)C carbon isotope ratios (delta(13)C-a time integrated measure of photosynthetic function) of needles in age classes II and IV were 3% (P < 0.01) and 2% (P < 0.05) more negative, respectively, in treated plants than in control plants. Exposure to enhanced UV-B radiation caused a 20% decrease in total biomass and a 4% (P < 0.05), 25% (P < 0.01), and 9% (P < 0.01) decrease in needle length of needles in age classes I, II, and IV, respectively. The observed decreases in delta(13)C, and F(v)/F(m) of the needles in the youngest needle age class suggest subtle damage to photosynthesis, although overall growth reductions were probably a result of decreased total leaf surface rather than decreased photosynthetic capacity. Needles of age class IV had lower light- and CO(2)-saturated maximum photosynthetic rates (39%), lower dark respiration (34%), lower light saturation points (37%), lower foliar nitrogen concentration (28%), and lower delta(13)C (14%) values than needles of age class I. Apparent quantum yield and F(v)/F(m) did not change with needle age. The observed changes in photosynthesis and foliage chemical composition with needle age are consistent with previous studies of coniferous trees and may represent adaptations of older needles to shaded conditions within the canopy.     

Effects of low night temperature and light on chlorophyll fluorescence of field-grown seedlings of Scots pine (Pinus sylvestris L.)

In vivo chlorophyll fluorescence kinetics was observed in naturally regenerated and planted Scots pine (Pinus sylvestris L.) seedlings growing at two adjacent sites in northern Sweden. Some seedlings were shaded from direct solar radiation and some were protected from frost at night. Air temperature and solar radiation were monitored. In vivo chlorophyll fluorescence kinetics of current-year needles was measured at both room temperature and 77 K. On clear days during August, variable fluorescence of photosystem II of needles from exposed, naturally regenerated seedlings was reduced by night frosts, whereas frost caused little change in fluorescence of needles from shaded seedlings. Overnight frost-protection reduced the inhibition of photosystem II by direct sunlight. Recovery from photoinhibitory damage occurred on cloudy days. In September, the decline of variable fluorescence of photosystem II was more pronounced in unshaded than in shaded seedlings, and coincided with frosts at night. In addition, the reduction in variable fluorescence was larger for planted seedlings than for naturally regenerated seedlings. It is suggested that the inhibition of photosystem II activity was caused by an interaction between low temperatures and light. Natural night frosts are proposed to increase the susceptibility to photoinhibition in the following day(s).     

Drought tolerance of clonal Malus determined from measurements of stomatal conductance and leaf water potential

We examined tolerance to soil drying in clonally propagated apple (Malus domestica Borkh.) rootstocks used to control shoot growth of grafted scions. We measured leaf conductance to water vapor (g(L)) and leaf water potential (Psi(L)) in a range of potted, greenhouse-grown rootstocks (M9, M26, M27, MM111, AR69-7, AR295-6, AR360-19, AR486-1 and AR628-2) as the water supply was gradually reduced. Irrespective of the amount of available water, rootstocks that promoted scion shoot growth (M26 and MM111) generally had higher g(L) and more negative Psi(L) than rootstocks that restricted scion shoot growth (M27 and M9). After about 37 days of reduced water supply, there were significant decreases in g(L) and Psi(L) in all rootstocks compared with well-watered controls. In all treatments, the slope of the relationship between log (g(L)) and Psi(L) was positive, except for rootstocks AR295-6, AR628-2 and AR486-1 in the severe-drought treatment, where the drought-induced change in the relationship suggests that rapid stomatal closure occurred when leaf water potentials fell below -2.0 MPa. This drought response was associated with increased root biomass production. Rootstock M26 showed little stomatal closure even when its water potential fell below -2.0 MPa, and there was no effect of drought on root biomass production. We conclude that differences among rootstocks in the way that g(L) and Psi(L) respond to drought reflect differences in the mechanisms whereby they tolerate soil drying. We suggest that these differences are related to differences among the rootstocks in their ability to control shoot growth.     

Some ecophysiological characteristics of artà (Calligonum comosum Ľ Hérit) in response to drought stress

Water deficit is an important environmental factor restricting plant growth and photosynthesis. The effect of water deprivation on leaf water status, photosynthetic gas exchange, chlorophyll content and fluorescence parameters of artà (Calligonum comosum) was studied. Five-month-old artà seedlings, grown in pots in the open air, were subjected to one of four drought treatments (i.e., mild, moderate, severe and extreme drought stress) and compared to control seedlings (normal watering regime). Results show that leaf water potential, net photosynthesis, stomatal conductance, transpiration, photosynthetic pigment content (chlorophyll a and b) decreased with increasing levels of drought stress. Inactivation of the photosynthetic apparatus was accompanied by changes in the fluorescence characteristics, providing evidence that reduction of photosynthetic rate could be attributed to non-stomatal limitations. Alterations imply changes in photochemical conversion efficiency of photosystem II by which plants could reduce water transpiration or protect their photosynthetic apparatus from destruction. These adaptations are discussed in relation to the strategies developed to grow drought-resistant artà seedlings in desert environments.