Intermittent low temperatures constrain spring recovery of photosynthesis in boreal Scots pine forests

During winter and early spring, evergreen boreal conifers are severely stressed because light energy cannot be used when photosynthesis is pre‐empted by low ambient temperatures. To study photosynthetic performance dynamics in a severe boreal climate, seasonal changes in photosynthetic pigments, chloroplast proteins and photochemical efficiency were studied in a Scots pine forest near Zotino, Central Siberia. In winter, downregulation of photosynthesis involved loss of chlorophylls, a twofold increase in xanthophyll cycle pigments and sustained high levels of the light stress‐induced zeaxanthin pigment. The highest levels of xanthophylls and zeaxanthin did not occur during the coldest winter period, but rather in April when light was increasing, indicating an increased capacity for thermal dissipation of excitation energy at that time. Concomitantly, in early spring the D1 protein of the photosystem II (PSII) reaction centre and the light‐harvesting complex of PSII dropped to their lowest annual levels. In April and May, recovery of PSII activity, chloroplast protein synthesis and rearrangements of pigments were observed as air temperatures increased above 0°C. Nevertheless, severe intermittent low‐temperature episodes during this period not only halted but actually reversed the physiological recovery. During these spring low‐temperature episodes, protective processes involved a complementary function of the PsbS and early light‐induced protein thylakoid proteins. Full recovery of photosynthesis did not occur until the end of May. Our results show that even after winter cold hardening, photosynthetic activity in evergreens responds opportunistically to environmental change throughout the cold season. Therefore, climate change effects potentially improve the sink capacity of boreal forests for atmospheric carbon. However, earlier photosynthesis in spring in response to warmer temperatures is strongly constrained by environmental variation, counteracting the positive effects of an early recovery process.     

Seasonal differences in photosynthesis between the C3 and C4 subspecies of Alloteropsis semialata are offset by frost and drought

The regional abundance of C₄ grasses is strongly controlled by temperature, however, the role of precipitation is less clear. Progress in elucidating the direct effects of photosynthetic pathway on these climate relationships is hindered by the significant genetic divergence between major C₃ and C₄ grass lineages. We addressed this problem by examining seasonal climate responses of photosynthesis in Alloteropsis semialata , a unique grass species with both C₃ and C₄ subspecies. Experimental manipulation of rainfall in a common garden in South Africa tested the hypotheses that: (1) photosynthesis is greater in the C₄ than C₃ subspecies under high summer temperatures, but this pattern is reversed at low winter temperatures; and (2) the photosynthetic advantage of C₄ plants is enhanced during drought events. Measurements of leaf gas exchange over 2 years showed a significant photosynthetic advantage for the C₄ subspecies under irrigated conditions from spring through autumn. However, the C₄ leaves were killed by winter frost, while photosynthesis continued in the C₃ plants. Unexpectedly, the C₄ subspecies also lost its photosynthetic advantage during natural drought events, despite greater water-use efficiency under irrigated conditions. This study highlights previously unrecognized roles for climatic extremes in determining the ecological success of C₃ and C₄ grasses.     

Recovery of photosynthetic capacity in <Emphasis Type="Italic">Vaccinium vitis</Emphasis>-<Emphasis Type="Italic">idaea</Emphasis> during mild spells in winter

Recent studies suggest that evergreen plants may maintain their photosynthetic capacity through the winter. Since mild winters are predicted to be more frequent in the future, the metabolic activity of plants is also likely to increase. The aim of the present study was to assess how various environmental factors, such as temperature, photoperiod and preceding frost, affect the recovery of photosynthesis during a mild spell in winter. The recovery of photosynthesis was studied in a series of growth chamber experiments where the overwintering of lingonberry (Vaccinium vitis-idaea) was interrupted by an intermittent warm spell of 1 week during different phases of winter. Rapid activation was observed in all the experiments during the first 3–4 days. No obvious effects of the phase of winter or photoperiod on the recovery of photosynthesis were observed, but a severe freezing treatment prior to the warm spell retarded the recovery significantly. Once recovered, however, lingonberry was able to maintain high rates of photosynthesis even at near-freezing temperatures, which prevail in their natural sub-nivean environment. The apparent quantum yield of photosynthesis remained high through the winter for lingonberry. This may prove advantageous for evergreen dwarf shrubs which overwinter in dim environments under snow.     

The after-effects of temperature and irradiance during early growth of winter oilseed rape (Brassica napus L. var. oleifera, cv. Górczański) seedlings on the progress of their cold acclimation

Experiments performed under controlled conditions showed that level of PPFD (photosynthetic photon flux density) during early seedlings growth (preceding cold acclimation at +2 °C) was not the key factor for the development of frost resistance. It did not modify the beneficial effects of prehardening (Rapacz 1997, in this issue) at moderately low (+12 °C) day temperature. Now I have shown that the increase of PPFD may replace to some extent prehardening in the development of frost resistance. It was particularly seen in non-prehardened plants, which had been grown under warm-day (+20 °C) conditions. Prehardening performed under controlled conditions, as well as seedlings growth under natural autumn conditions in the field, allowed to maintain a high net-photosynthesis rate at chilling temperatures. A net-photosynthesis rate during cold acclimation at +2 °C corresponded well with higher frost resistance. As a result, seedlings non subjected to prehardening and grown before cold acclimation under low PPFD acclimated better, if the cold treatment was applied only at nights (+20/2 °C day/night). Only under such conditions the photosynthetic rate was sufficiently high to allow plants to reach a higher level of frost resistance. All other plants acclimated better when they were exposed to the hardening temperature continuously during days and nights (+2/2 °C day/night).     

内蒙古半干旱草原区沙地植物群落光合特征的动态研究

沙蒿(Artemisia intramongolica)群落是半干旱草原地区沙地的重要植被类型,分别在植物的生长前期、中期、盛期和后期采用便携式光合测定仪和大型同化分析仪测定了沙蒿叶片和沙蒿群落的光合动态。单叶和群落的光合速率日进程类型随气候的不同而异,瞬时光合速率主要决定于光合有效辐射强度(PAR)。土壤干旱大大降低了单叶和群落的光合能力,晴天土壤湿润时气温和空气湿度控制着叶片的光合速率,午间大气湿度降低是光合午休的主要外因。叶片的蒸腾速率与气温呈显著线性相关,植物的光能和水分利用效率也主要取决于PAR和气温,随着PAR和气温的升高利用效率下降。沙蒿叶片光能利用效率在后期也能保持较高水平。沙蒿对土壤干旱和高温具有一定的适应性,在土壤湿润时能迅速提高光合速率,形成较大的生物量。但是沙蒿的蒸腾速率高,水分利用效率低。研究认为,沙蒿通过对土壤干旱和高温的忍耐机制而保持长时间较高的光能利用效率,并在土壤湿润时迅速提高光合能力和积累干物质来适应半干旱的沙地环境,而且依靠高蒸腾速率和强的水分吸收能力来竞争性抑制其他植物的生长。     

Effects of fern thickets on woodland development on landslides in Puerto Rico

Abstract. Thicket-forming ferns are common colonizers of disturbed habitats in the tropics, but little is known about their ecology. The effects of thickets formed by the fern Dicranopteris pectinata on tree seedlings on five landslides in the Luquillo Experimental Forest in northeastern Puerto Rico were both positive and negative. Soil moisture and total soil N were higher under fern thickets than in adjacent open areas and soil bulk density and soil surface temperatures were lower. Germination of seeds of the tree Cecropia schreberiana was higher for seeds sown under fern thickets than for those sown into adjacent open areas. Tree seedlings of Tabebuia hetero-phylla exhibited a threefold reduction in photosynthesis under ferns, probably resulting from a twelvefold reduction of photosynthetic photon flux density. Growth of Tabebuia seedlings was reduced under ferns but the distribution of seedlings of naturally occurring woody plants was not strongly correlated with the presence of fern thickets. Although fern thickets on low-nutrient landslide soils appear to facilitate germination, they inhibit growth of tree seedlings and may, therefore, delay forest development on landslides in Puerto Rico.     

Photosynthesis in frost-hardened and frost-stressed leaves of Hedera helix L.

Abstract The purpose of this study was to determine the respective extents to which winter reduction of photosynthetic capacity in ivy (Hedera helix L.) is caused by direct frost injury to the photosynthetic apparatus and by preceding protoplasmic changes connected with the acquisition of frost tolerance. Potted juvenile ivy plants were placed in the open under natural weather conditions whilst others were hardened under controlled conditions and subjected to the desired frost stress. Low non-freezing temperatures induced frost tolerance in ivy leaves down to about – 12°C (50% injury = TL₅₀) without impairing net photosynthetic rate as measured under standard conditions (20°C, light saturation, natural CO₂ level; = Standard-F_n. Only if the leaves froze (below ? 3°C to ?4°C) was a reversible inhibition of Standard-F_n observed. As long as the temperatures did not fall below approximately ?8°C the inhibition was small and Standard-F_n reached about 80–90% of the control. In this case the stomatal opening narrowed, giving a poorer supply of CO₂ to the mesophyll cells. Maximal frost tolerance (TL_5O?20°C to ?24°C) developed only with severe frosts below about ? 10°C. After such frosts, Standard-F_n was reduced to less than 20% of the control. The dependence of the rate of net photosynthesis on the internal CO₂ concentration showed a lower initial slope, thus indicating disturbances of chloroplast functions. However, neither in outdoor plants nor in those artificially frosted at – 20°C could there be found an appreciable inhibition of the electron transport capacity from H₂O to dichlorophenol indophenol or of ribulose bisphosphate carboxylase. If intact, severely frosted ivy plants were then held at higher temperatures (20/15°C), Standard-F_n recovered completely in approximately 10 d. Furthermore, following a frost period with temperatures down to ?12°C, mild weather caused a distinct improvement in Standard-F_n in outdoor plants, and there was no loss of maximum frost tolerance. Thus it can be concluded that the inhibition of Standard-F_n after severe frosts is not due to the development of maximal frost tolerance, but rather may be attributed to frost damage to the photosynthetic apparatus.     

Spring patterns of freezing resistance and photosynthesis of two leaf phenotypes of Hedera helix

Subdominant evergreen broad-leaved plants occurring in deciduous forests throughout temperate zones have only a short window of optimum photoassimilation in spring before canopy closure. Yet increasing photosynthetic and metabolic activity occurs concurrently with reductions in freezing resistance, resulting in vulnerability of plant tissues to late spring freezing events. Our goal was to document the temporal patterns of photosynthesis versus freezing resistance during spring in adult and juvenile leaf phenotypes of Hedera helix in Switzerland. Freezing resistances in all leaves were well below long-term minimum temperatures experienced at the study site, with adult leaf phenotypes in the forest canopy being more freezing resistant than juvenile leaves occurring closer to the ground. Reductions in freezing resistance were followed by increases in leaf photosynthetic capacities, which appeared synchronized among leaf phenotypes. Adult canopy leaves maintained a higher freezing resistance but lower photosynthetic capacity than juvenile leaves through the end of winter and into early spring. However, shortly after the cessation of freezing temperatures, adult leaves greatly increased their photosynthetic capacity relative to juvenile leaves, yet maintained freezing resistances sufficient to resist late spring freezing events. These patterns highlight the importance of the tradeoff in H. helix between exposure to potentially damaging cold temperatures in late spring and the need for high photosynthetic carbon gains before full canopy closure.     

Does the lack of vernalization requirement interfere with winter survival of oilseed rape plants?

Recent studies (Rapacz 1999) have shown that cultivars of spring-type oilseed rape are able to cold-acclimate to the level comparable with winter cultivars, but only after prehardening which results both in the increase of photosynthetic activity and in growth cessation. It is commonly known that under field conditions spring-type cultivars could not survive winter. Present studies were undertaken to explain the reasons for low winter hardiness of spring type rape plants. Six cultivars of spring and two of winter rape were sown in the open-air vegetation room at the end of August. The obtained results indicate that the degree of frost damage in spring-type plants increased in the course of winter and this increase was parallel to elongation of generative shoots observed after periods of warming. Each spring cultivar was completely killed by frost just after its generative shoot reached 15–20 cm, irrespective of its frost resistance level, determined previously under laboratory conditions. In the case of winter cultivars survival rate was consistent with laboratory-estimated frost resistance. It is suggested that spring rape could not survive winter because of its limited ability to prevent shoot elongation during winter at temperatures slightly above 0 °C. It was also found that less efficient photosynthetic electron transport in autumn was observed in these spring cultivars in which the elongation of generative shoots was observed already during the first warm break in winter.     

Morphophysiological monitoring of winter wheat in spring in the context of global climate warming

Data on morphophysiological monitoring of winter wheat (Triticum aestivum L.) cultivar Mironovskaya 808 grown in Hoagland and Arnon solution in a greenhouse and transferred to natural conditions in March–April 2004 with the mean daily temperature of 0.6 ± 0.7°C within the exposure period of 42 days are presented. Water content, dry weight of plants and their organs, frost hardiness of plants, degree of tissue damage by frost, CO₂ metabolism (photosynthesis and respiration), concentrations of sugars in tissues and proportions between different sugar forms, and activities of soluble and insoluble acid and alkaline phosphatases were monitored. Monitoring was carried out for three experimental variants simulating different microclimatic conditions in spring: after snow melting (experiment I), under ice crust (experiment II), and under snow cover (experiment III). Plants in experiments III and II demonstrated a higher water content in tissues, lower frost hardiness, higher rates of biomass loss, lower concentration of sugars and lower di-to monosaccharide ratio in tissues, and higher total invertase activity, particularly, cell wall-associated acid invertase activity. The dark respiration rates at 0°C did not significantly differ between experimental variants. The photosynthetic capacity at this measurement temperature was maintained in all experimental variants being most pronounced in experiment II with the most intense photoinhibition under natural conditions. Comparison of experiments III and II with experiment I is used to discuss the negative effect of changes in certain microclimatic indices associated with global warming and leading to plant exhaustion and death from frost in spring.     

Effects of artificial frost hardening and winter stress on net photosynthesis, photosynthetic electron transport and RuBP carboxylase activity in seedlings of Pinus silvestris

Net photosynthesis of seedlings of Pinus silvestris has been measured and compared with the activities of photosynthetic electron transport and extracted RuBP carboxylase. The effects of prolonged frost hardening (photoperiod 8 h, + 3°C) followed by winter stress at subzero temperatures were analysed. There was a parallel effect of frost hardening and winter stress on the photosynthetic properties of both intact seedlings and isolated chloroplast thylakoids. The activity of extracted RuBP carboxylase was less affected by the treatments. In relation to earlier works we conclude that the decay of net photosynthesis in winter climate is determined by the electron transport properties of the chloroplast thylakoids, i.e. by the pool sizes of photosynthetically active plastoquinone. The results of this work justify the definition of two phases in the response of conifers towards autumn and winter climates: I. Frost hardening occurs at temperatures slightly above zero and it does not affect the efficiency of photosynthesis as defined by the quantum yield at rate limiting light absorption. II. Winter stress occurs at subzero temperatures and it is characterized by a suppression of the photosynthetic efficiency as a result of damage within the photosynthetic apparatus.     

Early-spring gas exchange and uptake of deuterium-labelled water in the poikilohydric fernPolypodium virginianum

This study was carried out to determine if the desiccation-tolerant fernPolypodium virgimanum L. ecologically resembles lower plants by absorbing atmospheric water through its fronds and actively growing in early spring when the soil along cliff edges is still frozen. Three times between February and April, 1991,P. virginianum clonal mats were treated with deuterium-labelled water. Following each application, fronds were collected over several days and analyzed for the presence of deuterium. Two treatment groups plus a control were used: fronds were sprayed directly while covering the soil, or the roots were watered while protecting the fronds. The control mats were left untreated. Soil, air, and frond temperatures, plus photosynthesis and frond conductance were monitored throughout the study period. At subfreezing temperatures in February, no labelled water was taken up from the soil and no photosynthesis took place. Small amounts of label were absorbed from the soil in March during freeze-thaw cycles when rates of photosynthesis and stomatal conductance were both low. Large amounts of label were taken up from the soil in April when the soil was fully thawed and gas exchange was at normal seasonal levels. Label was not absorbed directly through the fronds when the plants were actively growing. Despite the desiccation tolerance ofP. virginianum, the timing and patterns of its water uptake and gas exchange in early spring resemble those found in higher vascular plants, not poikilohydric lower plants.     

A chamber for the simulation of radiation freezing of plants

Frost injury to plants can occur following episodic radiation frosts. In the UK this is particularly important to spring sown crops such as potatoes. Most laboratory based frost studies simulate freezing using either conductive or convective freezing chambers. Such frost tests do not simulate overnight freezing events adequately. A freezing chamber based on radiative cooling is described which mimics overnight radiative freezing. The chamber is rectangular in design (1 m × lm × 2 m high) with a radiative cooling plate at the top of the chamber cooled to -40°C to -45°C using HFC coolants, which acts as a cold black body. The sides of the chamber are also cooled to variable temperatures down to -5°C in order to prevent the chamber walls radiating to the plant material during testing. Using thermocouples to measure air temperature and plant temperature the chamber has been characterised to simulate the radiative cooling conditions found in the UK during autumn and spring. Exotherm detection upon plant freezing is simplified by virtue of the reduction in temperature fluctuation normally experienced at the plant surface during natural freezing. Radiation frosts and subsequent frost damage to potatoes have been recorded in the temperature range -4°C to –5°C. The equipment is recommended for studies of frost damage to plants normally caused by episodic radiation frost events.     

Effects of soil warming during spring on photosynthetic recovery in boreal Norway spruce stands

The effect of soil thawing and soil temperature on postwinter recovery of photosynthetic capacity was studied, during late spring and early summer, in Norway spruce stands in northern Sweden. Soil temperature was manipulated by means of buried heating cables. The warming treatment was applied to stands with low (natural) and high (fertilized) availability of nutrients. Soil thawing, expressed as water availability, was followed by means of sapflow in stems, and shoot water potentials. The recovery of photosynthetic capacity was assessed by measuring the rate of light-saturated photosynthesis (A_max), and maximum photochemical efficiency of photosystem II in detached shoots, and chlorophyll a fluorescence. Accumulation of starch reserves in the needles was followed as an independent indicator of photosynthetic performance in situ. Snowmelt and soil thawing occurred more than one month earlier in heated than in unheated plots. This was expressed both as sapflow and as differences in shoot water potential between treatments. During May, the rates of A_max were significantly higher on heated than on control plots. The effect of soil warming on A_max was, however, not reflected in chlorophyll fluorescence or needle starch content. The time course of the recovery of photosynthetic capacity was mainly controlled by mean air temperature and by the frequency of severe night frosts, and to a lesser extent by earlier soil thawing and higher soil temperatures.     

PHOTOSYNTHETIC SENSITIVITY TO TEMPERATURE IN POPULATIONS OF TWO C4 BOUTELOUA (POACEAE) SPECIES NATIVE TO DIFFERENT ALTITUDES

The relationships between photosynthesis, flowering, and growth temperatures were examined experimentally in four populations of the C₄ grass genus Bouteloua. Field-collected plants were grown under two temperature regimes, cool (20 C day/6 C night) and warm (30/16), representative of the extreme populations. Populations collected from the warm climates had significantly lower photosynthetic capacity when grown in the cool chamber relative to the warm chamber, while photosynthetic capacity in the cool climate populations did not differ between the growth conditions. Additionally, exposure to a 2-day cold temperature treatment (10/-2), representative of late-season frosts in high altitude sites, resulted in further reductions in photosynthesis in the warm climate plants, but not in the cool climate plants. This effect was greater for plants grown in the cool growth chamber. Flowering was reduced by 70% in the warm climate plants grown in the cool chamber, and was correlated with photosynthetic inhibition following the short-term cold temperature treatment. These results indicate that genetic differentiation for photosynthetic temperature sensitivity has occurred in the cool climate populations, and that long-term exposure to cool temperatures coupled with short-term relatively extreme low temperatures results in greater photosynthetic inhibition in nontolerant populations.     

不同土壤水分条件下中国沙棘雌雄株光合作用、蒸腾作用及水分利用效率特征

采用LI-6400便携式光合系统对野外不同生境(沟底、坡面)和田间不同土壤水分条件下中国沙棘雌雄株的净光合速率和蒸腾速率及水分利用效率特征进行了观测,结果表明,中国沙棘雌雄株在水分条件较好的生境中均表现较强的生活力,净光合速率和蒸腾速率主要受光照强度和大气温度的影响,雄株表现出更高的光合、蒸腾、水分利用效率;在水分条件较差的生境中,雌雄株均通过降低蒸腾和提高水分利用效率来适应逆境,净光合速率和蒸腾速率的影响因子增多,雄株的光合速率大幅度下降,雌株仍保持较高的光合速率,雌株的水分利用效率高于雄株.可见,在反映植物瞬时生理变化的性状方面,雌株对土壤水分减少表现出了更强的适应性,雄株对土壤水分变化表现出了更大的敏感性.     

羊草叶片气体交换参数对温度和土壤水分的响应

 采用生长箱控制的方法研究了羊草(Leymus chinensis)幼苗叶片光合参数对5个温度和5个水分梯度的响应和适应。结果表明：轻度、中度土壤干旱并没有限制羊草叶片的生长,对气体交换参数亦无显著影响,反映了羊草幼苗对土壤水分胁迫的较高耐性。叶片生物量以26 ℃时最大,其它依次为23 ℃、20 ℃、29 ℃和32 ℃。温度升高使气孔导度和蒸腾速率增加, 却使光合速率和水分利用效率降低。水分和温度对叶片生物量、光合速率、气孔导度和蒸腾速率存在显著的交互作用,表明高温加强了干旱对叶片生长和气体交换的影响, 降低了羊草对土壤干旱的适应能力。高温和干旱的交互作用将显著减少我国半干旱地区草原的羊草生产力。     

Photosynthetic capacity and light harvesting efficiency during the winter-to-spring transition in subalpine conifers

Some coniferous forest ecosystems undergo complete photosynthetic down-regulation in winter. The present study examined the influence of several environmental parameters on intrinsic, needle-level photosynthesis and photoprotection during the spring reactivation of photosynthesis in subalpine conifers. Maximal photosystem II (PSII) efficiency, photosynthetic capacity, and amounts of zeaxanthin and early light-inducible protein (Elip) family members were assessed in three subalpine conifer species over 3 years, and intensively during the 2003 winter-to-spring transition. During summers, maximal PSII efficiency remained high while intrinsic photosynthetic capacity varied depending on precipitation. During winters and the winter-to-spring transition, photosynthetic capacity and PSII efficiency were highly correlated and (during the spring transition) strongly influenced by air and soil temperature and liquid water availability. Decreases in the amount of Elip family members from winter through spring paralleled disengagement of sustained zeaxanthin-dependent photoprotection, although one of four anti-Elip antibody-reactive bands increased during spring. Intrinsic photosynthetic capacity and maximal PSII efficiency were highly responsive to day-to-day environmental changes during spring, indicating that multiple environmental signals are integrated to orchestrate the reactivation of photosynthesis from the inactive winter state to the active summer state.     

Microclimate above grass adversely affects spring growth of seedling snow gum (Eucalyptus pauciflora)

Growth of snow gum seedlings (Eucalyptus pauciflora Sieb. ex Spreng.) was studied in response to differences in microclimate caused by differential heat exchange between seedlings, grass and bare, moist soil during winter and spring. Seedlings were planted in a pasture either directly into grassy groundcover or in circular patches of bare soil of 30, 60 or 120 cm in diameter. There were no differences in maximum air temperatures at seedling leaf height between treatments. However, minimum air temperature increased by 2 °C with increase in patch diameter from 0 to 120 cm such that seedlings surrounded by grass experienced lower minimum temperatures with more frequent and more severe frosts than seedlings growing in large patches of bare soil. These small-scale differences in minimum temperature affected both photosynthetic and growth processes. Over winter, seedlings were photoinhibited, with depression in midday F_v/F_m linearly related to minimum temperatures. In spring, repeated frosts and lower minimum temperatures led to a delay in the recovery of F_v/F_m, a delay in bud-break, damage to elongating stems and developing leaves, lower rates of stem elongation, and ultimately a shorter growing season for seedlings in grass compared to those in bare soil patches. Thus, microclimate above grass adversely affects spring growth of juvenile Eucalyptus pauciflora and may account for much of the competitive inhibition of tree seedling growth by grass during spring.     

Canopy and leaf gas exchange of Haloxylon ammodendron under different soil moisture regimes

In order to reveal the drought resistance and adaptation of the C4 desert plant Haloxylon ammodendron under artificially controlled soil moisture regimes,representative plants were selected to measure canopy photosynthesis using canopy photosynthetic measurement system.The results showed that appropriate soil moisture significantly enhances the canopy and leaf photosynthetic capacity,and extremely high soil moisture is not conducive to the photosynthesis of H.ammodendron.