Stomatal and nonstomatal limitations of photosynthesis in relation to the drought and shade tolerance of tree species in open and understory environments

 Light saturated photosynthesis (A) in field saplings of shade tolerant, intermediate, and intolerant tree species was analyzed for stomatal and nonstomatal limitations to test differences between species and sun and shade phenotypes during drought. Throughout the study, photosynthesis was highest and mesophyll limitations of A (L_m) lowest in the intolerant species in both open and understory habitats. The shade tolerant species exhibited the only drought-related decreased A and increased L_m in the open, and the greatest drought-related decreased A and increased L_m in the understory. Few species exhibited significant habitat or drought-related differences in stomatal conductance to CO₂ (g_c), but even slight decreases in g_c during drought were associated with large increases in stomatal limitations to A (L_g). Combined changes in L_m and L_g resulted in increased relative stomatal limitation to A (l _g) in several species during drought. Nevertheless, the overall lack of stomatal closure allowed for nonstomatal limitations to play a major role in reduced A during drought. Higher leaf N was associated with shallower slope of the l _g versus g_c relationship, an indication of greater A capacity. Photosynthetic capacity tended to be greater in the intolerant species than the tolerant species, and it tended to decrease during drought primarily in the shade tolerant species in the understory. Findings in the literature suggest that carbon reduction reactions may be more susceptible to drought than photosynthetic light reactions. If so, reduced carbon reduction capacity of shade tolerant species or shade phenotypes may predispose them to drought conditions, which suggests a mechanism behind the well-recognized tradeoff between drought tolerance and shade tolerance of temperate tree species. Received: 20 October 1995 / Accepted: 20 February 1996     

Differential drought responses between saplings and adult trees in four co-occurring species of New England

Tree-ring characteristics in four species were examined to address whether co-occurring mature trees of different successional status respond differently to drought, and whether saplings of these species have a greater response to drought than mature trees. We examined saplings and mature trees of paper birch, yellow birch, red maple and sugar maple, which varied in successional status (shade-tolerance) and co-occurred at Harvard Forest, Petersham, Mass., USA. Three drought events in 1964–1966, 1981 and 1995 were identified using climate data. For mature trees, there was no significant interspecific difference in relative changes in ring-width index (RWI) during the 1964–1966 and 1995 drought events. However, the interspecific difference was significant in the 1981 drought event. Response function analysis for mature trees showed that the radial growth of sugar maple was mainly controlled by spring and summer precipitation, red maple by spring and summer precipitation and temperature, yellow birch by winter and summer precipitation, and spring and summer temperature, and paper birch by spring and summer precipitation and spring temperature. Saplings of sugar maple and yellow birch, but not red maple and paper birch, showed significant positive correlations between RWI and annual total precipitation. In the 1995 drought event, saplings and mature trees of red maple and paper birch differed significantly in drought responses, but this was not true in sugar maple and yellow birch. Our results do not support a generally greater response in saplings than in mature trees, nor an early- versus late successional difference in drought responses.     

Morphology and ecological significance of intra-annual radial cracks in living conifers

 Intra-annual radial cracks were studied on 294 cross-sections of Norway spruce sampled at two forest sites in the eastern Alps (Italy) and from seven isolated trees in the Jura region (Switzerland). Cracks were occasionally accompanied by traumatic resin canals in the wood that was formed after the cracking. Most of the cracks, however, were without such canals. Traumatic resin canals are not significantly more abundant in tree rings formed after cracking, and their occurrence is not related to the cracking. Cracks developed when the cambium was inactive. Water imbalances during the early spring, due to transpiration losses and inadequate moisture supply from very cold roots, are the likely cause of these cracks. Received: 21 February 1996 / Accepted: 14 June 1996     

Effects of seasonal rainfall on radial growth in two tropical tree species

 Seasonal drought may limit primary productivity in most of the tropics, but the determinants of tree growth are not well known. A 10-year study of the deciduous trees Cochlospermum vitifolium (Willd.) Spreng. (Cochlospermaceae) and Cnidoscolus spinosus Lundell (Euphorbiaceae) in southwestern México showed radial growth to be highly correlated (both r>0.85) only with precipitation during an interval of <2 months in the mid-wet season. Growth was not affected by total annual precipitation or by an early starting or late ending of the wet season, or by heavy rainfall in the dry season. Annual mean girth increments ranged from 0.03 to 3.31 cm and −0.1 to 2.01 cm, respectively. The best model for growth (r ²>0.85) was a linear combination of mid-summer precipitation (positive coefficient) and total precipitation over the previous 2 years (negative coefficient). Comparison with other species showed heterogeneous responses of wood production to climate variation, and suggests that the range of functional types of dry forest trees is still unknown. Received: 9 September 1996 / Accepted: 4 March 1997     

Seedling growth strategies and seed size effects in fourteen oak species native to different soil moisture habitats

 Seedling growth and morphology are thought to reflect evolutionary responses to habitat or influences of seed size. To test these hypotheses, we selected fourteen species of North American oaks differing in soil moisture habitat preference and seed size. Seedlings were grown for 1 – 2 years with abundant soil water and moderate soil nutrition in pots placed outdoors and in a common garden. Oak species native to xeric environments produced the smallest seedlings. Oaks from hydric soils had more shoot weight per unit of root weight and more height per unit of total plant weight than did mesic or xeric oaks. Essentially no differences in leaf area per unit of total plant weight were detected. Species with thinner and larger individual leaves tended to produce larger seedlings. Within species, seed size was generally unrelated to seedling growth, although results may have been complicated by uncontrolled genotypic variability. However, when species were compared, those with larger mean seed size produced larger seedlings. Root/shoot allometry, height growth and leaf thickness in the tested species may reflect evolutionary responses to soil moisture and flooding. Although seed size influenced seedling growth, no clear relationship between seed size and soil moisture habitat was found. Received: 26 March 1995 / Accepted: 30 November 1995     

Plant growth, leaf water potential, nitrogenase activity and nodule anatomy in Leucaena leucocephala as affected by water stress and nitrogen availability

 The effects of water stress and nitrogen availability on leaf water potential, nitrogenase activity, and growth was studied in a pot experiment with Leucaena leucocephala seedlings. Water stress was imposed on fertilized and unfertilized plants after inoculation with Rhizobium. Non-inoculated seedlings were used as control plants. Water stress lowered leaf water potential in all seedlings after 14 days of treatment. In inoculated seedlings, fertilized plants were more sensitive to water stress than unfertilized plants, as shown by a higher leaf water potential in plants of the latter treatment. Uninoculated and fertilized seedlings were most affected by water stress. This indicates that Rhizobium might increase stress tolerance in unfertilized seedlings at moderate water stress levels. The combined effects of water stress and applied fertilizers resulted in cessation of nitrogen fixation. Nitrogen fixation came to a complete stop after 22 days of water stress in fertilized seedlings. The different treatments were accompanied by anatomical changes of nodule structure. It is hypothesised that the leaf water potential may be used as an indicator to predict changes in nitrogen fixation in legume tree/shrub species during periods of water stress. Received: 21 October 1996 / Accepted: 12 May 1997     

Abrupt growth changes in Norway spruce and Yezo spruce near an industrial district in Hokkaido, Japan

 Increments in the radii of Norway spruce (Picea abies Karst.) and Yezo spruce (Picea jezoensis Carr.) trees that revealed symptoms of a decline in growth were analyzed by dendrochronological methods in an attempt to correlate past reductions in growth with their main causes. The trees were growing at different sites near the industrial district of Tomakomai, Hokkaido. A skeleton plot method was used to construct a series of pointer years that revealed the number of trees with a clear reduction in growth or recovery from such a reduction. An analysis of “abrupt growth changes” demonstrated that at least two periods of growth reduction were common to a large number of Norway spruce trees. The reduction events were related to the records of industrial activity near the forest and meteorological data. The growth reduction in the 1970s coincided with the start of operation of certain local factories, and its extent was related to the distance from the industrial region. By contrast, a reduction in growth in 1984 was detected at all the Norway spruce sites and the extent was approximately the same at all sites. This phenomenon was related to extreme drought conditions. Growth of Yezo spruce trees was less sensitive to industrial activity and to drought than that of Norway spruce. Thus, differences in response to air pollution and drought were observed between the two species. Received: 20 February 1996 / Accepted: 29 April 1996     

Radial growth responses to drought of Pinus sylvestris and Quercus pubescens in an inner‐Alpine dry valley

Question: Lower montane treeline ecotones such as the inner Alpine dry valleys are regarded as sensitive to climate change. In the dry Valais valley (Switzerland) the composition of the widespread, low altitude Pinus forests is shifting towards a mixed deciduous state. The sub‐boreal P. sylvestris shows high mortality rates, whereas the deciduous sub‐mediterranean Quercus pubescens is spreading. These species may act as early indicators of climate change. We evaluate this hypothesis by focusing on their differences in drought tolerance, which are hardly known, but are likely to be crucial in the current forest shift and also for future forest development. Methods: We used dendroecological methods to detect species‐specific patterns in the growth response to drought. The relationship between radial growth of 401 trees from 15 mixed stands and drought was analysed by calculating response functions using yearly tree‐ring indices and monthly drought indices. PCA was applied to the response ratios to discover spatial patterns of drought response. Results: A species‐specific response to moisture as well as a sub‐regional differentiation of the response patterns were found. While Quercus showed a response mainly to the conditions of the previous autumn and those of current spring, Pinus did not start responding before May, but showed responses throughout the whole summer. Quercus may restrict physiological activity to moist periods; growth of Pinus was much more dependent on prior growth. Conclusions: Given that the climate is changing towards (1) longer summer drought periods, (2) higher mean temperatures and (3) shifted seasonally of moisture availability, Quercus may benefit from adapting better to drier conditions. Pinus may increasingly face problems related to drought stress as it depends on summer moisture and has a smaller adaptive capacity due to its long‐lived photosynthetic tissue.     

Impact of long-term drought on xylem embolism and growth in Pinus halepensis Mill.

 The present study was carried out to elucidate the response mechanisms of 50-year-old Pinus halepensis Mill. trees to a long-term and severe drought. The amount of water available to trees was artificially restricted for 12 months by covering the soil with a plastic roof. Over the short term a direct and rapid impact of drought was evident on the water relations and gas exchanges of trees: as the soil dried out in the Spring, there was a concurrent decrease of predawn water potential; transpiration was strongly reduced by stomatal closure. Seasonal changes in the water volume fractions of twig and stem xylem were observed and interpreted as the result of cavitation and refilling in the xylem. When droughted trees recovered to a more favourable water status, refilling of embolized xylem was observed; twig predawn water potentials were still negative in the period when the embolism was reversed in the twig xylem. A few months after the removal of the covering, no differences in whole plant hydraulic resistance were observed between droughted and control trees. Needle and shoot elongation and stem radial growth were considerably reduced in droughted trees; no strategy of trees to allocate carbon preferentially to the stem conducting tissues was apparent throughout the experiment. An after-effect of the drought on growth was observed. Received: 4 August 1997 / Accepted: 1 October 1997     

Observed and projected impacts of climate on radial growth of three endangered conifers in northern Mexico indicate high vulnerability of drought-sensitive species from mesic habitats

A decline in productivity and radial growth in conifer forests from mesic areas has been associated with increased drought stress induced by climate warming. Nevertheless, studies showing how vulnerable tree species will be in response to forecasted warming conditions are scarce in such mesic habitats. Here we address this issue by analyzing how growth responds to drought and to observed and projected climate conditions in a conifer forest from northern Mexico, which is a hotspot of conifer diversity. We quantify the trends in radial growth (quantified as basal area increment, BAI) of three species (Abies durangensis, Picea chihuahuana, Cupressus lusitanica) using dendrochronology and a process-based model of tree growth. Growth decreased in A. durangensis and P. chihuahuana from the late 1980s onwards in response to warmer and drier conditions, whereas C. lusitanica growth showed very low sensitivity to precipitation and increased as temperature did. Winter-spring dry conditions adversely affected the growth of A. durangensis and P. chihuahuana. Our modeling approach anticipates growth reductions and an increase in the vulnerability of A. durangensis and the endangered P. chihuahuana against the warmer and more arid conditions predicted after the 2050s. Future warmer and drier climatic conditions could reduce the productivity and lead to growth decline of these mesic conifer forests triggering dieback episodes in highly drought-sensitive species as A. durangensis and P. chihuahuana.     

Effects of arbuscular mycorrhizal fungi and drought stress on growth and nutrient uptake of two wheat genotypes differing in drought resistance

 The effects of an arbuscular mycorrhizal (AM) fungus and drought stress on the growth, phosphorus, and micronutrient uptake of two wheat genotypes exhibiting differences in drought resistance were investigated. Plants were grown on a low P (4 mg kg^–1 soil) silty clay (Typic Xerochrept) soil-sand mix. Mycorrhizal infection was higher under well-watered than under dry soil conditions and the drought-resistant genotype CR057 had a higher mycorrhizal colonization than the drought-sensitive genotype CR006. Total and root dry matter yields and total root length were higher in mycorrhizal than in nonmycorrhizal plants of both genotypes. CR057 had higher total dry matter but not root dry matter than CR006 plants. The enhancement in total dry matter due to AM inoculation was 42 and 39% under well-watered and 35 and 45% under water-stressed for CR057 and and CR006, respectively. For both genotypes, the contents of P, Zn, Cu, Mn, and Fe were higher in mycorrhizal than in nonmycorrhizal plants and higher under well-watered than under dry soil conditions. The enhancement of P, Zn, Cu, Mn, and Fe uptake due to AM inoculation was more pronounced in CR006 than in CR057, particularly under water-stressed conditions. Thus CR006 benefitted from AM infection more than the CR057 under dry soil conditions, despite the fact that CR057 roots were highly infected. It appears that CR006 is more dependent on AM symbiosis than CR057. Accepted: 12 February 1997     

Contrasting effects of T cell growth factors on T cell responses to melanoma in vitro

 Determinants of T cell responses to tumor cells remain largely unknown. In the present study we have used long-term cultures of human melanoma cells and autologous peripheral blood lymphocytes to examine the influence of cytokines with T cell growth activity on the phenotype and cytotoxic and proliferative response of T cells to melanoma. It was found that addition of interleukin-4 (IL-4) inhibited the response of CD8⁺ T cells and promoted the response of the CD4 subset. IL-2 or IL-7 was effective in increasing melanoma-specific cytotoxic T lymphocyte (CTL) activity in cultures where CD8 T cells were predominant, whereas IL-4 followed by IL-2 was most effective in cultures where CD4 T cells predominated. IL-10 or IL-12 inhibited proliferation and CTL activity against melanoma in long-term cultures. The effects of IL-12 were reproduced in long-term cultures of T cells stimulated with mAb against CD3 and were shown to depend on prior exposure of T cells to IL-12 before IL-2. As yet unidentified factors, such as co-factor expression on melanoma, appear to be as important as exogenous cytokines in determining the nature of T cell responses to melanoma. These results suggest that analysis of responses in long-term culture may assist in defining the role of key cytokines and other determinants of immune responses to melanoma. Received: 4 June 1996 / Accepted: 12 November 1996     

山东蒙山森林冠层绿度与树干径向生长的关系

森林冠层绿度和树木年轮宽度是描述森林生长过程的重要指标,它们之间存在怎样的关系以及这种关系的稳定性如何目前还没有清晰的回答。森林冠层绿度通过遥感影像计算,在空间上连续,而树木年轮宽度是树木健康的综合指标,样点上具有代表性。森林冠层绿度和树木年轮宽度的关系的研究能增进对森林生长的多角度理解和森林生长状况的尺度转换。在山东蒙山地区采集了4个赤松(Pinus densiflora)林样点的树木年轮样本,获得了树木年轮宽度数据,分析了增强型植被指数(Enhanced Vegetation Index,EVI)与树木年轮宽度的关系。结果显示:1)对于健康森林,4月和6月的冠层绿度与树木年轮宽度存在因果关系;森林不健康时,两者关系较为复杂;2)其他月份冠层绿度与树干径向生长不存在因果关系,而是共同受其他环境因子,如气候因子的驱动;3)弱冠层绿度降低后5年内有显著的径向生长恢复,但是恢复年份少;强冠层绿度降低之前,树干径向生长已经开始降低,之后的5年内有着持续的径向生长降低。这些结果表明森林冠层绿度的降低并不能反映树干径向生长降低的开始,只有健康的森林冠层绿度和年轮宽度有相关关系。冠层绿度的降低对森林健康有强烈的影响,冠层绿度降低导致的径向生长的降低很难恢复。     

Changes in carotenoids, tocopherols and diterpenes during drought and recovery, and the biological significance of chlorophyll loss in Rosmarinus officinalis plants

Two-year-old rosemary (Rosmarinus officinalis L.) plants were subjected to severe stress by exposure to prolonged drought during a Mediterranean summer. Severely stressed plants recovered completely after the autumn rainfalls although the relative water content remained below 35% for 3 months and the chlorophyll content of leaves was reduced by up to 85% during the drought. In severe stress: (i) α-tocopherol increased 9-fold per g dry weight and 20-fold per unit of chlorophyll; (ii) lutein and β-carotene contents decreased on a dry-weight basis, but an 80% increase in lutein and constant levels of β-carotene were observed on a chlorophyll basis; (iii) there were transient and sustained increases in the de-epoxidation state of the xanthophyll cycle; and (iv) the highly oxidised abietane diterpene isorosmanol increased 8-fold as a result of the oxidation of carnosic acid. With the autumn rainfalls, water status, α-tocopherol and violaxanthin recovered first and the levels of photosynthetic pigments and abietane diterpenes increased later. The photoprotection conferred by the xanthophyll cycle and the antioxidant function of tocopherols, lutein and diterpenes may help to avoid irreversible damage in severe drought, making possible the recovery of functional membranes after the autumn rainfalls. Besides, chlorophyll loss reduces the amount of photons absorbed by leaves, which enhances the photoprotective and antioxidant capacity of leaves per amount of photons absorbed, since the ratios of xanthophylls, α-tocopherol and abietane diterpenes to chlorophyll increase. Received: 12 July 1999 / Accepted: 25 November 1999     

天山北坡前山带12种植物对干旱胁迫和复水的响应及抗旱性评价

【目的】探讨12种植物对干旱胁迫的动态生理适应性及抗旱能力,筛选适宜在天山北坡前山带进行植被恢复的灌木树种。【方法】以新疆天山北坡前山带种植的刺蔷薇、柠条锦鸡儿、文冠果、山杏、柽柳、梭梭、中亚沙棘、蒙古沙棘（‘新棘1号’、‘新棘2号’、‘新棘3号’、‘新棘4号’、‘新棘5号’）共12种植物为研究对象,观测灌水之后土壤水分的时空分布变化,分析干旱胁迫和复水处理对植物光合气体交换参数、抗逆生理生化指标、叶绿素荧光参数的影响,利用主成分分析法评价12种植物的抗旱能力。【结果】（1）在干旱胁迫发生前后,各植物立地土壤表层（0—20 cm）水分散失最快,中层（20—40 cm）次之,底层（40—60cm）水分散失最慢,而不同植物因其生长策略不同,通过改变其生长情况来适应干旱;（2）随着干旱胁迫的加剧,植物叶片光合生理指标净光合速率、气孔导度、蒸腾速率、胞间二氧化碳浓度、PSⅡ潜在活性、 PSⅡ最大光化学效率、电子传递速率和叶绿素含量降低,而抗逆生理指标超氧化物歧化酶活性、脯氨酸含量、丙二醛含量及非光化学猝灭系数升高,复水之后各指标都有一定程度的恢复,但均未达到初始水平;（3）主成分分析结果显示,净光合速率、叶绿素含量、电子传递效率和丙二醛含量等指标贡献率较大,梭梭、柽柳、刺蔷薇和柠条锦鸡儿抗旱性较强,中亚沙棘、‘新棘5号’、‘新棘1号’和山杏抗旱性次之,而‘新棘4号’、‘新棘3号’、文冠果和‘新棘2号’较弱。【结论】净光合速率、叶绿素含量、电子传递效率和丙二醛含量可作为评价各植物适应干旱胁迫的重要指标;新疆天山北坡前山带区域植被生态恢复过程中应优先考虑梭梭、柽柳、刺蔷薇及柠条锦鸡儿进行建植。     

Comparison of polyethylene glycol 3350 induced osmotic stress and soil drying for drought simulation in three woody species

 Drought simulation usually involves either soil drying or the use of an osmoticum, such as high molecular weight (>3000) polyethylene glycol (PEG). Although easy to apply, PEG absorption and toxicity remain a concern. This study compared the effects of soil drying and use of an osmoticum (PEG 3350). Osmotic stress and soil drought were applied to 5-month-old seedlings of jack pine (Pinus banksiana Lamb.) and black spruce [Picea mariana (Mill) B.S.P.] , which are both coniferous species from cold, boreal regions of North America, and flooded gum (Eucalyptus grandis W. Hill ex Maiden), a hardwood species growing in warmer, sub-tropical regions of Australia. Results showed that PEG 3350 was absorbed by roots, transported to shoots, and deposited on the leaves of both flooded gum and jack pine (but not black spruce). PEG lowered relative water content and damaged leaf tissues in both species, and also damaged stomata of flooded gum. Although 12 days of PEG-induced osmotic stress produced a decline in water potentials that was similiar to soil drying, it also caused significantly higher membrane injury and reduced net photosynthesis and stomatal conductance in leaves of all three species. Recovery of net photosynthesis and stomatal conductance in PEG-treated jack pine and black spruce was also slower after stress alleviation. Even a short exposure to PEG 3350 adversely affected seedlings compared to soil drought. These results confirmed that drought effects may vary, depending on the species and the method of stress induction. Received: 6 March 1996 / Accepted: 17 September 1996     

Provenance variation in morphology and leaflet anatomy of Parkia biglobosa and its relation to drought tolerance

 Measurements of morphological and anatomical characteristics made on 4-month-old seedlings from five provenances of Parkia biglobosa (Jacq.) Benth., grown in a tropical greenhouse under controlled conditions, showed significant differences between the northern provenances, which represented the Sudan savanna zone, and the southern provenances which represented the forest zone of the species distribution. Seedlings of provenances from the savanna zone showed more xerophytic characteristics than those from the forest zone. They were smaller in height, with a greater number of smaller leaflets per pinnae, a lower specific leaflet mass, smaller palisade and guard cells and a lower ratio of palisade:non-palisade tissue. There were also significant differences in the same characteristics, except palisade cell length, between provenances from the easterly and westerly extremes of the species range. Stomatal conductance measured after 2 weeks of droughting also showed that provenances from the savanna zone had higher values, indicating greater physiological activity and reduced stress, compared with provenances from the forest zone. Multivariate discriminant analysis of the morphological and anatomical characteristics showed that 98% of the seedlings were classified correctly with respect to provenance. The squared distances between provenances were closely related to the geographical distances between locations. Yet, not all variation in leaf anatomy could be explained by the difference in climatic conditions from north to south. Also drought tolerance was not always related to morphological and anatomical features. It is possible that clinal variation within the species from east to west may be interacting with climatic differences from north to south. Received: 2 June 1998 / Accepted: 29 June 1998