Leaf physiological versus morphological acclimation to high-light exposure at different stages of foliar development in oak

We investigated light acclimation in seedlings of the temperate oak Quercus petraea (Matt.) Liebl. and the co-occurring sub-Mediterranean oak Quercus pyrenaica Willd. Seedlings were raised in a greenhouse for 1 year in either 70 (HL) or 5.3% (LL) of ambient irradiance of full sunlight, and, in the following year, subsets of the LL-grown seedlings were transferred to HL either before leaf flushing (LL-HLBF plants) or after full leaf expansion (LL-HLAF plants). Gas exchange, chlorophyll a fluorescence, nitrogen fractions in photosynthetic components and leaf anatomy were examined in leaves of all seedlings 5 months after plants were moved from LL to HL. Differences between species in the acclimation of LL-grown plants to HL were minor. For LL-grown plants in HL, area-based photosynthetic capacity, maximum rate of carboxylation, maximum rate of electron transport and the effective photochemical quantum yield of photosystem II were comparable to those for plants grown solely in HL. A rapid change in nitrogen distribution among photosynthetic components was observed in LL-HLAF plants, which had the highest photosynthetic nitrogen-use efficiency. Increases in mesophyll thickness and dry mass per unit area governed leaf acclimation in LL-HLBF plants, which tended to have less nitrogen in photosynthetic components and a lower assimilation potential per unit of leaf mass or nitrogen than LL-HLAF plants. The data indicate that the phenological state of seedlings modified the acclimatory response of leaf attributes to increased irradiance. Morphological adaptation of leaves of LL-HLBF plants enhanced photosynthetic capacity per unit leaf area, but not per unit leaf dry mass, whereas substantial redistribution of nitrogen among photosynthetic components in leaves of LL-HLAF plants enhanced both mass- and area-based photosynthetic capacity.     

Photosynthetic acclimation to light changes in tropical monsoon forest woody species differing in adult stature

We studied morphological and physiological leaf and whole-plant features of seedlings of six late-successional woody species common in the Xishuangbanna lowland rain forest in southwest China. Study species differed in adult stature and shade tolerance and included the shrubs Lasianthus attenuatus Jack and Lasianthus hookeri C.B. Clarke ex Hook. f.; the sub-canopy species Barringtonia macrostachya (Jack) Kurz and Linociera insignis C.B. Clarke; the canopy tree Pometia tomentosa (Blume) Teijsm. & Binn.; and the emergent species Shorea chinensis (Wang Hsie) H. Zhu. After 1 year of growth in low light (4.5% full sun), seedlings were transferred to high light (24.5% full sun) to investigate acclimation responses of existing leaves to forest gap opening and to determine whether seedling capacity for acclimation is a limiting factor in its natural regeneration. Leaves of the shrub species are shade-adapted, as indicated by their low photosynthetic capacity, efficiency in using sunflecks, low stomatal density, low Chl a/b ratio and high spongy/palisade mesophyll ratio. The shrub species utilized sunflecks efficiently because they had a short photosynthetic induction time and low induction loss. In all species, transfer of seedlings to high light resulted in a substantial initial reduction in the dark-adapted quantum yield of photosystem II (variable chlorophyll fluorescence/maximum chlorophyll fluorescence; Fv/Fm) at midday. Predawn Fv/Fm of the taller species did not change greatly, but predawn Fv/Fm of the shrub species decreased significantly without complete recovery within 25 days of transfer to high light, indicating chronic photoinhibition and damage to the previously shade-adapted leaves. Maximum net photosynthetic rate and dark respiration of the four taller species increased considerably after transfer to high light, but not in the shrub species. Similar trends were observed for the number of newly formed leaves and relative height growth rate. We conclude that the shrubs L. hookeri and L. attenuatus have limited potential for developmental and physiological acclimation to high light, which explains their absence from forest gaps. Compared with the shrub species, the taller tree species, which are more likely to experience high light during their life span, showed a greater potential for light acclimation. Physiological differences among the four tree species were not consistent with differences in adult stature.     

Effects of the interaction between drought and shade on water relations, gas exchange and morphological traits in cork oak (Quercus suber L.) seedlings

The combined effect of drought and light on different physiological and biochemical traits was assessed in cork oak (Quercus suber L.) seedlings grown under two levels of light availability and submitted to a long-standing drought. Watering was withdrawn after germination and seedlings were allowed to dry to a water content of ca. 50% of field capacity. At this point, water-stressed seedlings were grown under moderate drought and two light regimes: high light (HL—50%) and low light (LL—2%). Soil water in control plants was kept close to field capacity (90–100%) for both light environments. Water-relations parameters derived from P–V curves, gas exchange and water status at predawn (Ψ_pd) were evaluated at twice during the experiment. Nitrogen and chlorophyll contents were determined in the same leaves used for the gas exchange measurements. In addition, maximum rate of carboxylation (V_cmax) and electronic transport (J_max) were derived from A–C_i curves in well-watered seedlings.

The variation on moisture availability during the experiment was the same under both light environments. In control plants, Ψ_pd was over −0.3 MPa at the two harvests, while stressed seedlings decreased to −0.9 MPa, with no differences between light treatments. Water stress decreased osmotic potentials at full (Ψπ₁₀₀) and zero turgor (Ψπ₀). The regressions between both potentials and Ψ_pd showed a higher intercept in shade grown seedlings. This fact will point out the higher osmoregulation capacity in sun seedlings whatever water availability.

Nitrogen investment on a per leaf mass (N_mass), chlorophyll content (Chl_mass) and SLA tended to show a typical pattern of sun-shade acclimation. Thus, the three parameters increased with shade. Only for N_mass there was a significant effect of watering, since water stress increased N_mass.

LL plants showed a lower photosynthetic capacity in terms of maximum net photosynthesis at saturating light (A_max), which was related to a decrease in V_cmax and J_max. Both parameters varied with specific leaf area (SLA) in a similar way. The low-light environment brought about a higher nitrogen investment in chlorophyll, while under high-light environment the investment was higher in carboxylation (V_cmax) and electronic transport (F_max).

Stomatal conductance to water vapour (g_wv) and A_max were lower in low-light seedlings independently of watering. In addition, there was a trend to keep higher intrinsic water use efficiency (IWUE) under high light environment. The increase of IWUE under water stress was higher in HL seedlings. This was as consequence of the steeper decline in g_wv as Ψ_pd decreased. The decrease of A_max with Ψ_pd occurred in a similar way in LL and HL seedlings. Thus, the HL seedlings tended to sustain a higher ability to increase IWUE than LL seedlings when they were submitted to the same water stress.     

Chlorophyll fluorescence as an indicator of frost hardiness in white spruce seedlings from different latitudes

This study examined the utility of variable chlorophyll fluorescence (F_var) to detect freezing damage in white spruce seedlings of four seedlots. Logistic regression analysis done for freezing tests in September showed that visible needle damage from freezing could be estimated by the F_var attributes F_o/I_ABS(r²=0.94), F_p(r²=0.98), F_v/F_m (r²=0.99), and F₁(r²=0.86). The regression curves indicated that for all four fluorescence attributes, inflection points occurred between 10 and 20% visible needle damage. The lack of a relationship between fluorescence attributes and visible seedling needle damage in October through December is because the minimum temperature (–18 and –24°C respectively) applied was insufficient to cause needle damage. Freezing-induced changes to F_var attributes can be detected which also result in photosynthetic rate decreases when no visible needle damage, and even electrolyte conductivity changes are evident. F_var attribute differences due to freezing can be resolved to the seedlot level. The Fvar curve feature manifested 5 seconds after dark-adapted seedlings have been exposed to light (F_5s) will estimate (r²=0.76) photosynthetic rate after freezing.     

Photosynthetic light acclimation in peach leaves: importance of changes in mass:area ratio, nitrogen concentration, and leaf nitrogen partitioning

Photosynthetic light acclimation of leaves can result from (i) changes in mass-based leaf nitrogen concentration, Nm, (ii) changes in leaf mass:area ratio, Ma, and (iii) partitioning of total leaf nitrogen among different pools of the photosynthetic machinery. We studied variations in Nm and Ma within the crowns of two peach (Prunus persica L. Batsch) trees grown in an orchard in Portugal, and one peach tree grown in an orchard in France. Each crown was digitized and a 3-D radiation transfer model was used to quantify the intra-crown variations in time-integrated leaf irradiance, . Nitrogen concentration, leaf mass:area ratio, chlorophyll concentration, and photosynthetic capacity were also measured on leaves sampled on five additional peach trees in the orchard in Portugal. The data were used to compute the coefficients of leaf nitrogen partitioning among carboxylation, bioenergetics, and light harvesting pools. Leaf mass:area ratio and area-based leaf nitrogen concentration, Na, were nonlinearly related to , and photosynthetic capacity was linearly related to Na. Photosynthetic light acclimation resulted mainly from changes in Ma and leaf nitrogen partitioning, and to a lesser extent from changes in Nm. This behavior contrasts with photosynthetic light acclimation observed in other tree species like walnut (Juglans regia L.) in which acclimation results primarily from changes in Ma.     

Influence of nutrient supply on shade-sun acclimation of Picea abies seedlings: effects on foliar morphology,photosynthetic performance and growth

Norway spruce seedlings (Picea abies Karst.) were grown in low light for one year, under conditions of adequate and limiting nutrition, then transferred to high light. Three months after transfer we measured photosynthesis, leaf nitrogen concentration, leaf chlorophyll concentration and leaf mass per area (LMA) of current-year and 1-year-old shoots; silhouette area ratio (SAR, the ratio of shoot silhouette area to projected needle area) was also measured in current-year shoots. At the foliage level, the effects of light and nutrient treatments differed markedly. Light availability during foliage expansion primarily affected LMA and SAR (morphological acclimation at the needle and shoot level, respectively). By contrast, nutrient supply in high light affected photosynthetic capacity per unit of leaf tissue (physiological acclimation at the cellular level) but did not affect LMA and SAR. The capacity for shade-sun acclimation in foliage formed before transfer to high light differed greatly from that of foliage formed following the transfer. The morphological inflexibility of mature needles (measured by LMA) limited their shade-sun acclimation potential. In contrast, at high nutrient supply, shoots that developed just after the change in photosynthetic photon flux density largely acclimated, both morphologically and physiologically, to the new light environment. The acclimation response of both current- and 1-year-old shoots was prevented by nutrient limitation. Analysis of growth at the whole-plant level largely confirmed the conclusions drawn at the shoot level. We conclude that nutrient shortage subsequent to the opening of a canopy gap may strongly limit the acclimation response of Norway spruce seedlings. Successful acclimation was largely related to the plant's ability to produce sun foliage and adjust whole-plant biomass allocation rapidly.     

Effect of cold storage treatments and transplanting stress on gas exchange, chlorophyll fluorescence and survival under water limiting conditions of Pinus radiata stock-types

The capacity of radiata pine seedlings to overcome planting shock in wet and dry conditions and their dependence on previous history was analysed by studying post-planting resumption of gas exchange and photochemical reactions, and survival 2 months later. Even under well-irrigated soil conditions, seedlings showed the effects of stress: gas exchange was reduced, but a clear difference between soil-plugged (PR) seedlings and bare-root (BR) seedlings was observed. Drought enhanced the severity of photosynthesis deprivation. Photochemical reactions, analysed by chlorophyll a fluorescence parameters, were not affected by planting shock in conditions of available soil water, but altered dramatically when drought stress was raised, suggesting structural damage of photosynthetic machinery. Despite the dramatic sensitivity of radiata pine to water availability, rewatering produced remarkable recovery, indicating good photosynthetic components repair capacity, which depended, however, on stock quality at the moment of planting. The ability of radiata pine to cope with drought in terms of post-planting performance depended on both storage conditions and water availability at the planting site. These findings provide information for tree physiologists and foresters as to how the management of radiata pine seedlings before planting can affect post-planting performance potential under wet or dry environmental conditions.     

Family differences in height growth and photosynthetic traits in three conifers

We investigated variation in height growth, gas exchange, chlorophyll fluorescence and leaf stable carbon isotope ratio among wind-pollinated progenies of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco var. glauca), ponderosa pine (Pinus ponderosa Dougl. ex Laws.) and western white pine (Pinus monticola Dougl. ex D. Don) from a small group of contiguous stands on the Priest River Experimental Forest in northern Idaho. Photosynthetic variables differed between height classes in the pines, but not in Douglas-fir. Among species and families, tall families of ponderosa pine regained photosynthetic capacity earliest in the spring and maintained it latest in the growing season. Tall families of western white pine had higher instantaneous water-use efficiencies and lower photosynthetic rates than short families on warm days in August.     

Biochemical acclimation patterns of Betula pendula and Pinus sylvestris seedlings to elevated carbon dioxide concentrations

Acclimation of photosynthesis to increasing atmospheric carbon dioxide concentration ([CO2]; 350 to 2,000 micromol mol-1) was followed in silver birch (Betula pendula Roth.) and Scots pine (Pinus sylvestris L.) seedlings for two years. Chlorophyll fluorescence and concentrations of Rubisco, chlorophyll, total soluble protein and nitrogen were monitored together with steady-state gas exchange at three CO2 concentrations (ambient [CO2] (345 +/- 20 micromol mol-1), the growth [CO2] and 1950 +/- 55 micromol mol-1). Rubisco and chlorophyll concentrations decreased in birch and Scots pine with increasing growth [CO2]. A nonlinear response was recorded for Rubisco and chlorophyll concentrations in birch, which was correlated with a significant decrease in specific leaf area. Nitrogen concentration decreased in birch leaves, but was unchanged in Scots pine needles. The species differed substantially in their steady-state CO2 exchange response to increasing growth [CO2]. The principal effect in birch was a significant nonlinear decrease in the steady-state gas exchange rate at the ambient [CO2], whereas in Scots pine the main effect was a significant increase in the steady-state gas exchange rate at the growth [CO2].     

Photosynthesis and root respiration of Pinus sylvestris and picea abies seedlings after different root freezing and storage temperatures

Photosynthetic performance and root respiration were measured for seedlings of Scots pine and Norway spruce under constant conditions in an open gas exchange system in the laboratory. Measurements were carried out after root exposure to ‐20, ‐5 and 0°C and subsequent longtime storage in darkness at +1 or +4°C. Stomatal conductance in relation to net photosynthetic rates was also investigated after the same treatment of seedlings. Root respiration was low for seedlings whose root system had been exposed to ‐20°C, Scots pine showing lower rates than Norway spruce. This was probably an indication of root damage. At least for one provenance of Scots pine, respiration rates were higher for seedlings stored at +1 than at +4°C. Photosynthetic performance was also lowest for seedlings whose roots had been exposed to +20°C compared to higher temperatures, the difference being more clear‐cut for Norway spruce than for Scots pine. Storage at +1 gave slightly higher photosynthetic rates than at +4°C. There was a close relation between stomatal conductance measured on individual needles and photosynthetic performance measured on the whole seedling.     

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)]     

Acclimation of leaf characteristics of Fagus species to previous-year and current-year solar irradiances

To examine the effects of different solar irradiances on leaf characteristics at the leaf primordium and expansion stages, we shaded parts of branches in the upper canopies of two adult beech trees, Fagus crenata Blume and Fagus japonica Maxim., for 4 years. The treatments during the leaf primordium and leaf expansion stages, respectively, were: (1) high light and high light (H, control), (2) high light and low light (HL), (3) low light and low light (LL), and (4) low light and high light (LH). Both number of cell layers in palisade tissue and individual leaf area were affected by the previous-year irradiance, whereas cell length of palisade tissue was larger in LH leaves than in LL leaves, suggesting determination by current-year irradiance. Lamina chlorophyll/nitrogen ratio was higher in HL and LL leaves than in LH leaves, suggesting determination by current-year irradiance. Diurnal minimum values of leaf water potential measured under sunlit conditions were lower in H and LH leaves than in HL and LL leaves. Effective osmotic adjustment was found in H and LH leaves, suggesting that leaf water relations were affected by current-year irradiance. Net photosynthetic rate and stomatal conductance measured under sunlight conditions were higher in H and LH leaves than in HL and LL leaves. Thus, effects of current-year irradiance had a greater effect on leaf-area-based daily carbon gain than previous-year irradiance.     

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.     

Seasonal acclimation of photosystem II in Pinus sylvestris. I. Estimating the rate constants of sustained thermal energy dissipation and photochemistry

Acclimation of the partitioning of absorbed light energy in Photosystem II (PSII) between photochemical and non-photochemical processes includes short-term adjustments that are rapidly reversed in the dark and seasonal acclimation processes that are unaffected by dark acclimation. Thus, by using dark-acclimated leaves to study the seasonal acclimation of PSII, the confounding effect of short-term adjustments is eliminated. The maximum quantum yield of photochemistry, estimated by chlorophyll fluorescence analysis as F(v)/F(m), where F(v) = (F(m) - F(o)), and F(m) and F(o) are maximum and minimum chlorophyll fluorescence, respectively, has been widely used to follow the seasonal acclimation of PSII, because it is measured in dark-acclimated leaves. Seasonal changes in F(v)/F(m) can be caused by adjustments in either the photochemical capacity in PSII, or the capacity of thermal dissipation in PSII, or both. However, there is a lack of chlorophyll fluorescence parameters that can distinguish between these processes. In this study, we introduce two new parameters: the rate constants of sustained thermal energy dissipation (k(NPQ)) and of photochemistry (k(P)). We estimated k(NPQ) and k(P) from dark-acclimated F(o) and F(m) measured during spring recovery of photosynthesis in Scots pine (Pinus sylvestris L.) trees. We suggest that k(NPQ) and k(P) be used to study the mechanisms underlying the observed seasonal acclimation in PSII, because these parameters provide quantitative data that complement and extend F(v)/F(m) measurements.     

Seasonal acclimation of photosystem II in Pinus sylvestris. II. Using the rate constants of sustained thermal energy dissipation and photochemistry to study the effect of the light environment

Photosynthesis in evergreen conifers is characterized by down-regulation in autumn and rapid up-regulation in spring. This seasonal pattern is largely driven by temperature, but the light environment also plays a role. In overwintering Scots pine (Pinus sylvestris L.) trees, PSII is less down-regulated and recovers faster from winter stress in shaded needles than in needles exposed to full sunlight. Because the effect of light on the seasonal acclimation of PSII has not been quantitatively studied under field conditions, we used the rate constants for sustained thermal energy dissipation and photochemistry to investigate the dynamics and kinetics of the seasonal acclimation of PSII in needles exposed to different light environments. We monitored chlorophyll fluorescence and needle pigment concentration during the winter and spring in Scots pine seedlings growing in the field in different shading treatments, and within the crowns of mature trees. The results indicated that differences in acclimation of PSII in overwintering Scots pine among needles exposed to different light environments can be chiefly attributed to sustained thermal dissipation. We also present field evidence that zeaxanthin-facilitated thermal dissipation and aggregation of thylakoid membrane proteins are key mechanisms in the regulation of sustained thermal dissipation in Scots pine trees in the field.     

Seasonal changes in chlorophyll fluorescence of white spruce seedlings from different latitudes in relation to gas exchange and winter storability

Variable chlorophyll fluorescence (F_var) was investigated as a tool in detection of distinct seasonal physiological changes in 1+0 intact white spruce seedlings. The loss of the characteristic F_var peak (Fp) between 0.8 and 1.0 s after illumination of dark adapted seedlings is an indication of regulation of photosynthetic activity in August. The peak represents excess photochemical water-splitting of photosystem II. We interpret its loss as a physiological indicator of the process of dormancy induction. Three dimensional (i.e. X[0-300 s], Y[rfu], Z[time of year] axis) F_var curves of non-stressed seedlings measured over 300 s followed a three phase change over the growing season. In actively growing seedlings, the portion of the Kautsky induction curve between 60 and 300 s was 0.4 relative fluorescence units (rfu) in northern (i.e. >56° latitude) seedlots and 0.6 in the southern (i.e. <56° latitude) range seedlot from August until early September. About mid-September curve features between 60 and 300 s decreased sharply to approximately zero (rhu) by October. Freeze test data indicated seedlings became frost hardy during this time. The third, or inactive phase was seen as flat line from 40 to 100 s. The portion of the curve after 100 s was responsive to short term temperature changes. White spruce seedlots of northern and southern B.C. latitudes having curve fluorescence peak (F_p) values at about 1 s of 0.6, and 0.8 (rhu) respectively, plus curve minimum (F_min) values at about 60 s which do not decrease further over a 2–3 week period represent stock which can safely be lifted for cold storage. The F_var attribute at 5 s after the actinic light is turned on (F_5s) correlates well with net photosynthesis (r² =0.61) during the growing season.     

Growth and ecophysiology of seedlings of Podocarpus falcatus in plantations of exotic species and in a natural montane forest in Ethiopia

The potential role of exotic tree plantations in facilitating successional processes on degraded areas was evaluated in southern Ethiopia by comparing seedling characteristics, transpiration and photo-synthetic performance of Podocarpus falcatus seedlings in Eucalyptus plantation, Pinus plantation, adjacent natural forest and clear-felled plantation site. P. falcatus seedlings exhibited differences in architecture between Eucalyptus and Pinus plantations. They had higher leaf area, shorter internode length and greater number of lateral branches in Eucalyptus plantation. At similar vapor pressure deficit (VPD), P. falcatus transpired much less than E. saligna, especially at higher VPDs. Analysis of fluorescence parameters in the leaves showed no significant differences in the level of dark-adapted and light-adapted fluorescence yield (Fv /Fm and ΔF/Fm, respectively), electron transport rate (ETR) and nonphotochemical quenching (NPQ) among seedlings grown inside plantations and adjacent natural forest, indicating similar photosynthetic performance. Nevertheless, there was evidence of photoinhibition in P. falcatus in the clear-felled site which had low fluorescence yield but high values of NPQ as protection from photoamage. The light response curves of ETR, NPQ and ΔF/F m , showed similar light saturation behavior among the seedlings grown inside plantations and natural forest and suggested a sequence of light-adapted to shade-adapted behavior in Natural forest > Eucalyptus plantation > Pinus plantation. The results show the structural flexibility, better water-use and adaptability of P. falcatus in its use of the understory environment of plantation species.     

Responses to temperature and shade in Abies alba seedlings from diverse provenances

Abstract

Intraspecific variability in responses to temperature and shade was studied at Champenoux, north-eastern France, with seedlings from five Polish provenances of silver fir (Abies alba Mill.). Acclimation of photosynthesis to temperature was investigated in seedlings exposed to 10, 25 and 35°C in a climate chamber for 1 week. During two growth seasons, a population of seedlings was grown in the nursery under four different irradiance regimens: 100, 48, 18 and 8% of natural irradiance. Maximum carboxylation rate (V _cmax), maximum light driven electron flow (J _max) and maximum net carbon dioxide assimilation rate (A _max) measured at 25°C increased with population altitude. One week of exposure to 35°C caused discoloration and massive needle shedding. After 2 years’ acclimation to different levels of irradiance, a significant interprovenance variability was evidenced in growth, total biomass, biomass allocation and photosynthetic performance. This study provided evidence for the existence of functional variation among the examined provenances.     

3个与杉木混植木荷种源幼苗对光照响应的差异

目的]探寻不同木荷种源混交林生产力差异的光合机制。[方法]以福建建瓯、江西信丰和浙江龙泉3个木荷代表性种源1年生和2年生幼苗为试验材料,在与杉木混植条件下,设置100%(L0)、50%(L1)和25%(L2)3种光照的光环境,研究3个木荷种源幼苗的生长性状、光合生理及叶绿素荧光特性等对不同光照响应的差异。[结果]1年生和2年生木荷幼苗的苗高、地径和干质量均表现出显著的种源差异,福建建瓯种源幼苗生长优于浙江龙泉种源和江西信丰种源。随着遮阳强度的提高,1年生和2年生2个发育阶段,福建建瓯种源幼苗比叶面积和叶绿素含量等明显增加,叶绿素a/b值、光饱和点、暗呼吸速率和Fo值等均显著降低,表现出较高的形态和生理塑性;相同遮阳处理下福建建瓯种源幼苗总叶面积、叶绿素含量和Fv/Fm值等指标显著高于浙江龙泉种源和江西信丰种源,而叶绿素a/b值、光饱和点、光补偿点和暗呼吸速率则明显低于两种源;长期遮阳和混植状态下福建建瓯种源幼苗可通过较高的生物量分配塑性,增加根系在土壤中分布来响应邻株竞争,进而促进整株干物质的积累;各光环境下,3个木荷种源2年生幼苗Fo、Fm、Fv/Fm和Fv/Fo等叶绿素荧光参数均较1年生幼苗明显升高,但各光环境间差异不显著。[结论]不同木荷种源混交林生产力差异与其对光照的塑性反应能力有关,并随幼苗年龄而变化。     

模拟酸雨对两种针叶植物气体交换和叶绿素荧光特性的影响

以马尾松(Pinus massoniana)和杉木(Cunninghamia lanceolata)为试验材料,通过盆栽的方法,研究了重度酸雨处理、中度酸雨处理和酸雨对照处理对这两种针叶植物叶片叶绿素相对含量、气体交换参数以及叶绿素荧光特性等的影响.结果表明:在pH4.0时,酸雨胁迫对马尾松和杉木幼苗的地径生长有一定的促进;在不同酸雨处理下,马尾松幼苗的最大净光合速率从大到小依次是pH5.6>pH4.0>pH2.5;杉木幼苗的最大净光合速率从大到小依次是pH2.5>pH4.0>pH5.6,说明这2种植物幼苗的光合能力受到影响;Fv/Fm和Fv/Fo随着酸雨酸度的增加而减小,变化趋势是pH5.6>pH4.0>pH2.5,说明在pH 2.5的酸雨胁迫下,两种针叶植物幼苗叶片受到胁迫,叶绿素分子捕获激发能的效率和PSⅡ潜在活性有所降低.