Exploring the growth response of Norway spruce (Picea abies) along a small-scale gradient of soil water supply

The climate-growth response of specific sites and species is one of the main research subjects in classic tree ring studies. Traditional sampling approaches therefore aim at maximizing the climate signal of the analyzed tree ring series, which is typically achieved by focusing on dominant trees or on sites located in particularly temperature or moisture limited environments. However, there is increasing evidence that these selective sampling strategies cannot yield chronologies that are representative for entire populations. One promising approach to gain a deeper understanding of forest dynamics and climate-growth responsiveness is the analysis of climate signal ranges among trees. This individualistic approach requires random sampling and the integration of information on small-scale heterogeneities in site and tree characteristics. Here, we analyze the climate-growth response of 144 Norway spruce trees (Picea abies Karst.) on difference levels of data aggregation. The aim of our study is to investigate the relevance of small-scale heterogeneities in site conditions, particularly in soil water supply, for the detected climate-growth signal. We identify soil water supply and site characteristics, which indirectly modify the water availability for trees, as dominating growth factors across scales. The driest sites show the strongest climate-growth reaction, while the growth response of wetter sites is weak or even insignificant. Therefore, we conclude that integrating small-scale information on site characteristics, particularly on soil water supply, can help to gain a deeper understanding of species specific growth limitations.     

Carbon dioxide assimilation and growth of red spruce (Picea rubens Sarg.) seedlings in response to ozone,precipitation chemistry,and soil type

Summary The influence of ozone, mist chemistry, rain chemistry, and soil type on CO₂ assimilation and growth of red spruce (Picea rubens Sarg.) seedlings was investigated over a 4-month period under controlled laboratory and glasshouse conditions. Growth was evaluated through interval estimates of aboveground relative growth rates (RGR) and the partitioning of biomass components at harvest to root, stem, and needle fractions. Precipitation chemistry treatments and O₃ exposure dynamics were based on reported characteristics of air chemistry and/or deposition in high-elevation forests of eastern North America. The two soils were collected from Camels Hump in the Green Mountains of Vermont and Acadia National Park on the Maine coast. Soil from Acadia had higher organic content, higher levels of extractable base cations, and lower levels of extractable aluminum and heavy metals. The only treatment variables that consistently influenced the growth of P. rubens were soil type and rain chemistry. In comparison with seedlings grown in soil from Acadia National Park, those grown in Camels Hump soil had significantly less needle (27%), stem (33%), and root (26%) biomass at harvest and statistically lower aboveground RGR within 2 months after initiation of the treatments. Seedlings grown in Camels Hump soil had significantly higher levels of aluminum (6.5X), copper (1.4X), and nickel (2.7X) in new needle tissue. The only influence of precipitation chemistry on the growth of P. rubens was a pattern of greater root and shoot biomass in seedlings experiencing the more acidic rain treatments. Interactive effects among the main treatment variables (e.g., acidic mist and O₃, acidic rain and soil type) on seedling growth were not notable. Rates of CO₂ assimilation and transpiration on a per gram needle dry weight basis [mol·g^-1·s^-1] were not influenced by any of the main treatment variables or their interaction. Because neither soil type nor precipitation chemistry influenced the efficiency of CO₂ assimilation per gram dry weight of needle tissue, the physiological mechanism underlying the growth response of P. rubens is attributed to a change in either whole-plant allocation of carbon resources or a direct toxic effect in the rhizosphere on root growth.     

降雨和土壤湿度对贵州旱田土壤N2O释放的影响

以南方亚热带代表性旱田土壤-贵州玉米-油菜轮作田、大豆-冬小麦轮作田和休耕地为观测对象,研究土壤N2O释放通量季节变化与降雨和土壤湿度的关系,同时,采用DNDC模型定量探讨了未来降雨量变化对土壤N2O释放的潜在影响,结果表明,降雨与N2O释放峰间存在明显的驱动-响应关系,N2O释放通量与降雨量和土壤湿度间存在正相关性,模型检验结果表明,夏秋季土壤N2O释放通量与降雨量变化呈正相关,而降雨量的大幅度增加或下降将引起冬春季土壤N2O释放通量的微弱下降。     

全球变暖对太行山植被生产力及土壤水分的影响

将相同的自然植被用Lysimeter从高海拔下移至低海拔,温度升高2℃,同时设置平均降水,增加10％降水,增加20％降水,减少10％降水和减少20％降水5个降水处理,模拟全球变暖带来的温度升高和降水变化对植被生产力和土壤水分的影响,两年的野外实验表明,温度升高造成生态适应性差的野古草（Arundinelia hirta)生产力显著下降,致使整个测试群落生产力降低,低海拔实验点生产力显著低于高海拔实验点,温度升高对铁杆（Artemisia sacrorum)和黄背草（Themeda japonica)的影响较小,太行山区的植被对降水的变化反映,降水增加使植被的生产力水平显著提高,其中降水增加20％的处理生产力比平均降水处理的生产力增加22％,增加降水处理的土壤含水量显著高于平均降水和降水减少的实验处理,由于植被的减少,温度升高的实验点从第二年开始土壤水分较高。     

苜蓿与沙打旺苗期生长和水分利用对土壤水分变化的反应

通过室内生长箱内盆栽实验,比较了苜蓿和沙打旺苗期的根冠生长和水分利用对种土壤水分环境变化的响应和差异.结果表明,充分供水下苜蓿和沙打旺苗期生物量和蒸腾效率均最高,苜蓿均显著高于沙打旺.土壤水分减少后苜蓿苗期生物量和蒸腾效率下降幅度均大于沙打旺.从低水到阶段低水处理后土壤水分逐渐降低和降低后再复水到低水处理,苜蓿和沙打旺的生物量分别较持续低水处理显著减少47.8%和27.9%.旱后复水后苜蓿根冠比和单位根量耗水量较显著增加,蒸腾效率显著下降; 沙打旺根冠比显著下降,单位根量耗水量和蒸腾效率变化不显著.     

Willow water uptake and shoot extension growth in response to nutrient and moisture on a clay landfill cap soil

Extension growth of willow (Salix viminalis L.) and changes in soil water were measured in lysimeters containing clay and sandy loam soils with different amendment and watering treatments. No water uptake was found below 0.3 m in the nutritionally poor unamended clay; amendment with organic matter to 0.4 m depth resulted in water extraction down to 0.5 m depth whereas in the sandy loam, there was greater extraction from all depths down to 0.6 m. With water stress, wilting of plants occurred when the volumetric soil water content at 0.1 m was about 31% in the clay and 22% in the sandy loam. Compared with shoots on plants in the amended clay, those in the unamended treatment showed reduced extension growth, little increase in stem basal area (SBA) and a small shoot leaf area, resulting from a reduced number of leaves shoot⁻¹ and a small average area leaf⁻¹. Water stress also reduced shoot extension growth, SBA gain and the leaf area on extension growth. Shoot growth rates were significantly correlated with air temperature and base temperatures between 2.0 and 7.6 °C were indicated for the different treatments. These studies have helped to explain some of the large treatment effects described previously on biomass production and plant leaf area.     

Historical precipitation predictably alters the shape and magnitude of microbial functional response to soil moisture

Soil moisture constrains the activity of decomposer soil microorganisms, and in turn the rate at which soil carbon returns to the atmosphere. While increases in soil moisture are generally associated with increased microbial activity, historical climate may constrain current microbial responses to moisture. However, it is not known if variation in the shape and magnitude of microbial functional responses to soil moisture can be predicted from historical climate at regional scales. To address this problem, we measured soil enzyme activity at 12 sites across a broad climate gradient spanning 442–887 mm mean annual precipitation. Measurements were made eight times over 21 months to maximize sampling during different moisture conditions. We then fit saturating functions of enzyme activity to soil moisture and extracted half saturation and maximum activity parameter values from model fits. We found that 50% of the variation in maximum activity parameters across sites could be predicted by 30‐year mean annual precipitation, an indicator of historical climate, and that the effect is independent of variation in temperature, soil texture, or soil carbon concentration. Based on this finding, we suggest that variation in the shape and magnitude of soil microbial response to soil moisture due to historical climate may be remarkably predictable at regional scales, and this approach may extend to other systems. If historical contingencies on microbial activities prove to be persistent in the face of environmental change, this approach also provides a framework for incorporating historical climate effects into biogeochemical models simulating future global change scenarios.     

祁连山西部青海云杉径向生长对气候因子的响应

利用树木径向生长监测仪(Dendrometer)和自动气象站在祁连山西部对8棵青海云杉的径向生长及环境因子进行了连续监测研究。结果表明: Gompertz函数拟合结果显示,2018—2020年青海云杉的径向生长分别开始于4月19日、4月17日和4月10日,在日平均气温超过5.5 ℃时径向生长开始;青海云杉生长结束时间分别为8月17日、8月21日和7月19日,生长结束时间与生长季末期降水量有关。研究区青海云杉径向生长量受干旱抑制强烈,其中与7月的日均温(负相关)和日降水量(正相关)相关性最高,与生长季初期(5月)的日降水量的相关关系存在年际间差异。     

Changes in the radial growth of Picea crassifolia and its driving factors in the mid-western Qilian Mountains,Northwest China since 1851 C.E

Examining the growth of trees in response to environmental factors is essential for evaluating the stability of forest ecosystems. In this study, using tree ring data obtained from 18 sites and climate diagnostic methods. we investigated relationships between the radial growth of Picea crassifolia in the mid-western Qilian Mountains and local climate/sea surface temperatures (SSTs) since 1851 C.E. The results revealed that the radial growth of P. crassifolia showed significant upward trends during three time periods (1884–1906, 1929–1946, and 1964–1983) and significant downward trends in a further three periods (1907–1928, 1947–1963, and 1984–1995). Variations in the growth of P. crassifolia showed a significant negative correlation with temperature in June and a positive correlation with precipitation from July in the previous year to June in the current year as a response to climate change. We also found that large-scale anomalies could influence the radial growth of P. crassifolia, which was reflected in results showing that extremely high radial growth is related to El Niño patterns in the central-eastern equatorial Pacific, whereas extremely low growth is related to anomalously warmer SSTs in the southern Indian Ocean. Notably, we found that the extremely low growth of P. crassifolia in response to SST was more stable than that of extremely high growth. Furthermore, on the basis of qualitative methodology, we established that years characterized by extremely high/low growth were largely/little influenced by the time window and threshold values that were selected when determining the positive/negative pointer years. Our results confirm the validity of using the relationships between extremely low growth and SSTs to predict forest dynamics.     

Moisture-driven changes in the sensitivity of the radial growth of Picea crassifolia to temperature,northeastern Tibetan Plateau

Precipitation is one of the most important climate factors controlling tree growth, yet it is not fully understood how changes in precipitation affect the relationship between growth and temperature. On the northeastern edge of the Tibetan Plateau, nine tree-ring chronologies of Picea crassifolia were developed along a precipitation gradient from semi-arid (mean annual precipitation, 255 mm) to semi-humid (710 mm). We analyze the growth-climate relationships along this precipitation gradient and assess whether these associations are regulated by local precipitation. From 1960 to 2014, temperature increased significantly while precipitation remained stable at the nine sampling sites. The radial growth of P. crassifolia decreased at the semi-arid sites but increased at the semi-humid sites. Growth-temperature relationships gradually changed from negative to positive along the precipitation gradient (from dry to wet sites), particularly during summer. The moist P. crassifolia sites are also characterized by positive correlations with the Palmer Drought Severity Index. The temporal growth-temperature relationships varied significantly among the different spruce sites over the last five decades. Although temperature remains the main factor controlling the growth of P. crassifolia, local precipitation variability is becoming increasingly important. Our findings indicate that considering species distribution areas supports the analyses of the impact of climate change on tree growth.     

黄土丘陵区不同土地利用的土壤水分灰色关联度

运用灰色关联度分析法,分析了黄土丘陵区的延安燕沟不同土地利用方式下土壤水分的垂直变化和月动态.结果表明:梯田各层土壤水分变化态势相似程度较高,即梯田土壤水分的垂直波动较小;不同土地利用方式下均是表层(0-30cm)和中层(30-100cm)土壤水分的灰关联度较大,即土壤水分的变化发展态势较一致,但由于不同利用方式的影响,土壤水分变化态势的相似程度为梯田〉白羊草〉刺槐〉沙棘,除白羊草地外,其他土地利用方式下表层(0-30cm)与深层(100-200)的灰关联度最小,土壤水分变化差异较大.从各月土壤水分灰关联度来看,沙棘除10月份外,其他各月土壤水分变化态势的相似程度较高;白羊草地正好相反,整个生长季土壤水分的变化波动很大;梯田和刺槐林地居中,但各月土壤水分变化态势的相似程度仍然存在差异,表现为雨季前各月土壤水分变化较为一致,雨季后的9、10月份与4月份相比差异较大.说明不同的土地利用方式对土壤水分的垂直变化和月动态变化均产生不同影响. 态势的相似程度为梯田》白羊草》刺槐》沙棘,除白羊草地外,其他土地利用方式下表层(0-30cm)与深层(100-200)的灰关联度最小,土壤水分变化差异较大.从各月土壤水分灰关联度 看,沙棘除10月份外,其他各月土壤水分变化态势的相似程度较高;白羊草地正好相反,整个生长季土壤水分的变化波动很大;梯田和刺槐林地居中,但各月土壤水分变化态势的相似程度仍然存在差异,表现为雨季前各月土壤水分变化较为一致,雨季后的9、10月份与4月份相比差异较大.说明不同的土地利用方式对土壤水分的垂直变化和月动态变化均产生不同影响. 态势的相似程度为梯田》白羊草》刺槐》沙棘,除白羊草地外,其他土地利用方式下表层(0-30cm)与深层(100-200)的灰关联度最小,土壤水分变化差异较大.从各月土壤水分灰关联度 看,沙棘除10月份外,其     

On-line screening of soil VOCs exchange responses to moisture, temperature and root presence

The exchanges of volatile organic compounds (VOCs) between soils and the atmosphere are poorly known. We investigated VOC exchange rates and how they were influenced by soil moisture, temperature and the presence of plant roots in a Mediterranean forest soil. We measured VOC exchange rates along a soil moisture gradient (5%–12.5%–20%–27.5% v/v) and a temperature gradient (10°C–15°C–25°C–35°C) using PTR-MS. Monoterpenes were identified with GC-MS. Soils were a sink rather than a source of VOCs in both soil moisture and temperature treatments (−2.16 ± 0.35 nmol m⁻² s⁻¹ and −4.90 ± 1.24 nmol m⁻² s⁻¹ respectively). Most compounds observed were oxygenated VOCs like alcohols, aldehydes and ketones and aromatic hydrocarbons. Other volatiles such as acetic acid and ethyl acetate were also observed. All those compounds had very low exchange rates (maximum uptake rates from −0.8 nmol m⁻² s⁻¹ to −0.6 nmol m⁻² s⁻¹ for methanol and acetic acid). Monoterpene exchange ranged only from −0.004 nmol m⁻² s⁻¹ to 0.004 nmol m⁻² s⁻¹ and limonene and α-pinene were the most abundant compounds. Increasing soil moisture resulted in higher soil sink activity possibly due to increases in microbial VOCs uptake activity. No general pattern of response was found in the temperature gradient for total VOCs. Roots decreased the emission of many compounds under increasing soil moisture and under increasing soil temperature. While our results showed that emission of some soil VOCs might be enhanced by the increases in soil temperature and that the uptake of most soil VOCs uptake might be reduced by the decreases of soil water availability, the low exchange rates measured indicated that soil-atmosphere VOC exchange in this system are unlikely to play an important role in atmospheric chemistry. Electronic Supplementary Material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Effect of soil moisture stress on legume-rhizobium symbiosis in soybeans

Summary In a pot culture experiment, the influence of soil moisture stress at different physiological stages of soybean, cv. Hark, on nodulation, symbiosis and nitrogen accumulation was studied. Moisture stress reduced leghemoglobin content of root nodules and nitrogen uptake by plants. It had no effect on number of bacteroids. Stress at mid bloom and rapid pod filling stages reduced yield and seed protein content. However, these parameters were not affected by stress at nodule initiation and early flowering stages, though, flower initiation and maturity of the plant were delayed. Moisture stress at any stage did not alter nitrogen status of roots.     

Intra-annual radial growth of Schrenk spruce (Picea schrenkiana Fisch. et Mey) and its response to climate on the northern slopes of the Tianshan Mountains

Schrenk spruce (Picea schrenkiana Fisch. et Mey.) is widely distributed in the Tianshan Mountains. In this study, four Schrenk spruce trees were continuously monitored with dendrometers from 27 April to 30 September 2014 on the northern slopes of the Tianshan Mountains in northwest China. The goal of this monitoring study was to determine the main growing season of Schrenk spruce and to analyze intra-annual radial growth variability and its relation to daily meteorological factors. Our studies have shown that the critical growing season of Schrenk spruce is from late May to late July and that the rapid growth stage is from mid-June to early July. Meanwhile, in the growing season, changes in the radial growth of Schrenk spruce were negatively correlated with daily temperature, evaporation, sunshine hours and vapor pressure deficit (VPD), and were positively correlated with precipitation and relative humidity (RH). The correlation coefficient between radial growth and RH can be as high as 0.750 (Pearson, p < 0.0001, n = 60). Dates in which precipitation occurred corresponded to periods of rapid growth. The results of the climate-growth analysis show that changes in radial growth reflect the effect of water stress on tree growth, whether or not the changes are positively or negatively correlated with the above climatic factors. This indicates that moisture plays a major role in the growth of Schrenk spruce. We suggest that precipitation between late May to late June is a limiting factor for radial growth of Schrenk spruce on the northern slopes of the Tianshan Mountains.     

黄土丘陵区植被恢复的土壤碳水效应

黄土高原大规模植被恢复显著影响了这一区域土壤水分和有机碳(SOC),从而影响其承载的土壤水源涵养和固碳服务。明确深层土壤水分和有机碳对植被恢复的响应特征是当前黄土高原地区生态水文与生态系统服务研究的一个重要科学问题,其中植被类型以及生长年限是这一过程的重要影响因素。然而,目前关于深层土壤有机碳和土壤水分对植被恢复的响应及二者关系的研究较少。通过对陕北典型黄土丘陵区不同植被类型和生长年限下0—5 m土壤水分与有机碳的监测,分析了深层土壤水分和有机碳对植被恢复的响应及其特征。研究发现:(1)植被恢复后0—5 m土层均出现水分亏缺,土壤水分亏缺在表层1 m最低,2—3 m最高;对于不同恢复方式,林地土壤水分亏缺在恢复至21—30a时显著高于前一阶段(11—20a),而在恢复31a后水分开始恢复,而灌木、草地土壤水分亏缺程度则随恢复年限延长不断增加。(2)林地、灌木、草地0—5 m平均土壤有机碳含量为1.97、1.77、1.72 g/kg;林地土壤固碳量随恢复年限的增加而增加,并且在恢复20a时固碳量与对照农田相比出现净增;灌木土壤固碳量随恢复年限先增加后降低;草地土壤固碳量则随退耕年限增加呈下降趋势并且低于对照农田。(3)表层0—1 m土壤水分随恢复年限增加变化不显著,深层土壤水分则随恢复年限增加显著降低;相比而言,随恢复年限增加,土壤有机碳随年限的变化在各层土壤中均不显著。深层土壤水分与土壤有机碳呈现显著的正相关,且土壤有机碳的增加速率低于土壤水分,研究认为,深层土壤固碳与土壤水分关系密切,且深层土壤固碳需要充足水分参与。深层土壤水分亏缺可能限制植被细根的发展,使深层土壤有机碳输入减少。     

Seasonal precipitation and continentality drive bimodal growth in Mediterranean forests

Tree phenology is sensitive to climate warming and changes in seasonal precipitation. Long xylogenesis records are scarce, thus limiting our ability to analyse how radial growth responds to climate variability. Alternatively, process-based growth models can be used to simulate intra-annual growth dynamics and to better understand why growth bimodality varies along temperature and precipitation gradients. We used the Vaganov-Shashkin (VS) growth model to analyse the main climatic drivers of growth bimodality in eight trees and shrubs conifers (four pines and four junipers) across Spain. We selected eleven sites with different continentality degree and spring/autumn precipitation ratios since we expected to find pronounced bimodal growth in less continental sites with spring and autumn precipitation peaks. The VS model successfully simulated annual growth rates at all sites as a function of daily temperature and soil moisture data. Bimodal growth patterns clustered into less continental sites showing low spring/autumn precipitation ratios. This finding agrees with observed climate-growth associations showing that growth was enhanced by wet-cool winter-to-spring conditions, but also by wet autumn conditions in the most bimodal sites. We observed a stronger growth bimodality in pines compared to junipers. We discuss the spatial variability of climate drivers in bimodality growth pattern and how increasing continentality and shifts in seasonal precipitation could affect growth patterns. Bimodality could be an advantageous response to overcome summer drought in Mediterranean forests. The ability of some species to reactivate growth during autumn might determine their capacity to withstand increasing summer aridity.     

Leaf attributes and tree growth in a tropical dry forest

Questions: How are leaf attributes and relative growth rate (RGR) of the dominant tree species of tropical deciduous forest (TDF) affected by seasonal changes in soil moisture content (SMC)? What is the relationship of functional attributes with each other? Can leaf attributes singly or in combination predict the growth rate of tree species of TDF? Location: Sonebhadra district of Uttar Pradesh, India. Methods: Eight leaf attributes, specific leaf area (SLA); leaf carbon concentration (LCC); leaf nitrogen concentration (LNC); leaf phosphorus concentration (LPC); chlorophyll concentration (Chl), mass‐based stomatal conductance (Gs_mass); mass based photosynthetic rate (A_mass); intrinsic water use efficiency (WUEi); and relative growth rate (RGR), of six dominant tree species of a dry tropical forest on four sites were analysed for species, site and season effects over a 2‐year period. Step‐wise multiple regression was performed for predicting RGR from mean values of SMC and leaf attributes. Path analysis was used to determine which leaf attributes influence RGR directly and which indirectly. Results: Species differed significantly in terms of all leaf attributes and RGR. The response of species varied across sites and seasons. The attributes were positively interrelated, except for WUEi, which was negatively related to all other attributes. The positive correlation was strongest between Gs_mass and A_mass and the negative correlation was strongest between Gs_mass and WUEi. Differences in RGR due to site were not significant when soil moisture was controlled, but differences due to season remained significant. The attributes showed plasticity across moisture gradients, which differed among attributes and species. Gs_mass was the most plastic attribute. Among the six species, Terminalia tomentosa exhibited the greatest plasticity in six functional attributes. In the step‐wise multiple regression, A_mass, SLA and Chl among leaf attributes and SMC among environmental factors influenced the RGR of tree species. Path analysis indicated the importance of SLA, LNC, Chl and A_mass in determining RGR. Conclusion: A _mass, SMC, SLA and Chl in combination can be used to predict RGR but could explain only three‐quarters of the variability in RGR, indicating that other traits/factors, not studied here, are also important in modulating growth of tropical trees. RGR of tree species in the dry tropical environment is determined by soil moisture, whereas the response of mature trees of different species is modulated by alterations in key functional attributes such as SLA, LNC and Chl.     

Effect of seed size on seedling growth response to elevated CO2 in Picea abies and Picea rubens

Several previous studies have observed that species and individuals with large seeds respond more positively to elevated CO (2) than those with small seeds. We explored the reasons for this pattern by examining the relationship between seed size and CO (2) response in Picea abies and P. rubens using growth analysis. The large seeded species (P. abies) responded more positively to elevated CO (2) than the small seeded species (P. rubens). At the intraspecific level, P. abies individuals from large seeds responded more positively to elevated CO (2) than individuals from small seeds, however, there was no significant intraspecific variation in CO (2) response in P. rubens. The greater CO (2) response of plants from large seeds was not simply the result of a larger starting capital compounded at the same rate as in plants from small seeds. Elevated CO (2) increased relative growth rate to a greater extent in individuals from large seeds. This effect appears to be related to differences in time of establishment, source to sink ratio and nutrient availability with seed size. These results are significant not only in understanding the potential effect of rising atmospheric CO (2) concentrations on plant populations, but also in understanding the factors affecting plant success at current atmospheric CO (2) levels due to the elevation of CO (2) within the litter layer that occurs at many germination sites.     

Growth,development and yield of pawpaw (Carica papaya L.) ‘Homestead selection’ in response to soil moisture stress

Summary The effects of four soil water potential regimes under greenhouse conditions on growth, development, and yield of ‘Homestead selection’ pawpaw were investigated. The objectives were to determine the critical soil water potential regime and the moisture sensitive stages of growth of pawpaw. Repeated cycles of stress to −6.0 bar soil water potential imposed as from vegetative phase prevented fruit formation by constantly causing flower abscission. The stressed plants were stunted in size. The −2.0 bar soil water potential value was considered the critical level for normal growth and reproductive development of pawpaw. The mid-vegetative, flowering and fruit enlargement phases were moisture sensitive.