Separating water‐potential induced swelling and shrinking from measured radial stem variations reveals a cambial growth and osmotic concentration signal

The quantification of cambial growth over short time periods has been hampered by problems to discern between growth and the swelling and shrinking of a tree stem. This paper presents a model, which separates cambial growth and reversible water‐potential induced diurnal changes from simultaneously measured whole stem and xylem radial variations, from field‐measured Scots pine trees in Finland. The modelled growth, which includes osmotic concentration changes, was compared with (direct) dendrometer measurements and microcore samples. In addition, the relationship of modelled growth and dendrometer measurements to environmental factors was analysed. The results showed that the water‐potential induced changes of tree radius were successfully separated from stem growth. Daily growth predicted by the model exhibited a high correlation with the modelled daily changes of osmotic concentration in phloem, and a temperature dependency in early summer. Late‐summer growth saw higher dependency on water availability and temperature. Evaluation of the model against dendrometer measurements showed that the latter masked a true environmental signal in stem growth due to water‐potential induced changes. The model provides better understanding of radial growth physiology and offers potential to examine growth dynamics and changes due to osmotic concentration, and how the environment affects growth.     

杉木径向生长动态及其对季节性干旱的响应

为探讨杉木径向变化的季节动态及其气候响应特征,利用径向生长仪连续2年(2016—2017年)监测了江西中部杉木的径向变化过程,分析了径向变化的日动态、季节动态规律及其与气候因子的相关性。结果表明: 杉木日径向昼夜变化呈白天收缩、夜间膨胀的格局;2017年径向生长开始时间比2016年提前一个月,但旱季持续的水分亏缺使生长季也早一个月结束;在主要生长季内(4—9月),无论湿季与旱季,径向增长量与降雨、相对湿度呈显著正相关,与光合有效辐射、饱和水汽压差呈显著负相关,而水分亏缺量的气候相关性与径向增长量相反;旱季严重缺水时土壤含水量对径向变化的影响显著增强。水分条件始终是影响杉木径向变化的关键因素,夏季干旱时可通过提高土壤含水量等有效途径促进杉木径向生长。     

Remote sensing tracks daily radial wood growth of evergreen needleleaf trees

Relationships between gross primary productivity (GPP) and the remotely sensed photochemical reflectance index (PRI) suggest that time series of foliar PRI may provide insight into climate change effects on carbon cycling. However, because a large fraction of carbon assimilated via GPP is quickly returned to the atmosphere via respiration, we ask a critical question—can PRI time series provide information about longer term gains in aboveground carbon stocks? Here we study the suitability of PRI time series to understand intra‐annual stem‐growth dynamics at one of the world's largest terrestrial carbon pools—the boreal forest. We hypothesized that PRI time series can be used to determine the onset (hypothesis 1) and cessation (hypothesis 2) of radial growth and enable tracking of intra‐annual tree growth dynamics (hypothesis 3). Tree‐level measurements were collected in 2018 and 2019 to link highly temporally resolved PRI observations unambiguously with information on daily radial tree growth collected via point dendrometers. We show that the seasonal onset of photosynthetic activity as determined by PRI time series was significantly earlier (p < .05) than the onset of radial tree growth determined from the point dendrometer time series which does not support our first hypothesis. In contrast, seasonal decline of photosynthetic activity and cessation of radial tree growth was not significantly different (p > .05) when derived from PRI and dendrometer time series, respectively, supporting our second hypothesis. Mixed‐effects modeling results supported our third hypothesis by showing that the PRI was a statistically significant (p < .0001) predictor of intra‐annual radial tree growth dynamics, and tracked these daily radial tree‐growth dynamics in remarkable detail with conditional and marginal coefficients of determination of 0.48 and 0.96 (for 2018) and 0.43 and 0.98 (for 2019), respectively. Our findings suggest that PRI could provide novel insights into nuances of carbon cycling dynamics by alleviating important uncertainties associated with intra‐annual vegetation response to climate change.     

Patterns of daily stem growth in different tree species in a warm-temperate forest in northern China

Radial growth in trees responds to environmental changes in various ways ranging from immediate to hysteretic responses. However, species-specific tree radial growth patterns and their responses to short-term weather changes are not fully understood. Here, the daily stem radial changes (SRCs) in four common tree species, linden (Tilia mongolica), birch (Betula dahurica), oak (Quercus wutaishanica) and larch (Larix principis-rupprechtii), were monitored with high-resolution point dendrometers during the main growing seasons in 2017–2019 on Dongling Mountain, northern China. The SRC was differentiated into tree water deficit-induced stem shrinkage (TWD) and growth-induced irreversible stem expansion (GRO) to evaluate species-specific responses to weather variables and short-term drought events. We found that the TWD and GRO of the four species were significantly different. The TWD was influenced primarily by the vapor pressure deficit (VPD), whereas the GRO was influenced primarily by precipitation (P). In linden and birch, a larger proportion of the GRO occurred at higher air temperature (T_mean) and VPD values; in contrast, the range of these changes was lower in oak and larch. With the increased durations of drought periods, oak and larch experienced large and rapid increases in TWD, whereas birch and linden showed small and slow increases. These results indicate that oak and larch would be sensitive to warmer and drier weather conditions predicted for the future, while linden and birch would have a conservative growth strategy. Our results provide further insights into the physiology of these four tree species and allow us to better predict the growth response of forest dynamics under climate change.     

Experimental analysis of the role of water and carbon in tree stem diameter variations

The variations of stem diameter as they can be accurately measured by Linear Variable Differential Transformer (LVDT) reflect the addition of four components: irreversible radial growth, reversible living-cell dehydration/rehydration, thermal expansion and contraction, and expansion of dead conducting elements due to the increase and relaxation of internal tensions. The correct interpretation of LVDT signals, with respect to the practical applications, should make an exact distinction between these four components. This paper describes a set of two experiments with potted hybrid walnut trees. Double girdling, water stress, and duration of the day versus night periods were used in the phytotron as experimental factors to induce variations of the carbon and water status of plant tissues. The latter were assessed, respectively, by water potential and transpiration, and by local stem respiration and carbohydrate content. The results are interpreted in terms of carbon or water limitation effects on stem diameter variations where radial growth and tissue elasticity could be distinguished. Moreover, they suggest no or very low involvement of CO2 originating from a distance, i.e. carried by the transpirational flux of xylem sap, in the total stem CO2 efflux rate.     

祁连山东部青杄年内径向生长动态对气候的响应

树木径向生长对气候因子的响应是树轮气候学的基础。在我国西北地区,虽然已有大量的树轮-气候响应研究,但是响应分析多基于数理统计结果,缺乏对树木生长过程的理解。基于此,于2013年到2015年在祁连山东部吐鲁沟国家森林公园内通过Dendrometer连续监测了树轮气候重建中常用树种青杄的径向生长,通过平均值法提取2013到2015年生长季内不同时间尺度(1天、7天、10天)的平均径向生长量及对应时间段的平均气候状况。不同时间尺度径向生长量和气候因子的相关分析结果表明,时间尺度的延长削弱了树干水分昼夜变化的干扰,但水分仍然是青杄径向生长的限制因子。为了进一步分析青杄径向生长与气候关系随时间的变化情况,以31天为窗口将日径向生长量与气候要素每隔一天进行滑动相关,结果显示:5、7月青杄的径向生长与降水的关系稳定,都呈显著正相关,但是6月降水的年际变率较大,导致树木可利用水分的变幅也较大,因而树木径向生长与6月水分的响应关系不稳定,存在较大的年际差异,而这可能是一些树轮-气候响应研究中轮宽与6月降水关系不显著的原因。     

Carbon allocation between tree root growth and root respiration in boreal pine forest

Soil respiration, i.e. respiration by mycorrhizal roots and by heterotrophic organisms decomposing above- and below-ground litters, is a major component in ecosystem carbon (C) balances. For decades, the paradigm has been that the biomass of fine roots of trees turns over several times a year, which together with large inputs of above-ground litter leaves little room for the contribution from root respiration. Here, we combine the results of a recent tree girdling experiment with the C budget of the classic Swedish Coniferous Forest (SWECON) project, in which root growth and turnover were estimated to be high. We observe that such a high rate of root turnover requires an unlikely high C use efficiency for root growth, and is not consistent with the 1:1 relation between root: heterotrophic respiration obtained in the girdling experiment. Our analysis suggests that 75% of the C allocated to roots is respired, while 25% is used for growth, and hence that root growth and turnover were grossly overestimated in the SWECON study.     

Simulated and predicted responses of tree stem radial growth to climate change—A case study in semi-arid north central China

Precise knowledge how tree growth will respond to future climate change is essential for the adapted management of forest ecosystems. By conducting sensitivity tests, tree-ring process-based cambial growth models can provide an innovative way to better understand wood formation under different climate change scenarios. As a case study in semi-arid north central China, we used artificially increased or decreased daily climatic data as input to the Vaganov-Shashkin dynamic growth model to investigate the response of wood formation to climatic change. By calibrating the tree-ring model using daily climate data over the period 1951–2010, we found that 81% of radial growth was driven by soil moisture, while 13% of growth was controlled by temperature. During the main growing season June–August, significant differences in the integral growth rate occurred after changing precipitation by ± 30% or by decreasing temperature by 3.0 °C (p < 0.05). However, increasing temperature showed only modest effects on tree radial growth rate. During the past 60 years, a significant advancement of the starting dates of growth was detected, whereby non-significant variability was found for the ending dates of growth. Contemporaneously, the effect of previous winter temperature (previous December to current January) on cambial growth initiation declined after 1980. Significant differences in the growth onset dates only occurred when temperature was reduced by 4.5 °C or increased by 5.5 °C. Moreover, both the onset and ending dates of growth in the study region were more sensitive to cooling rather than to warming. If temperature will increase by 2°C and precipitation will increase by 30% at the end of this century as predicted by some Earth system models, tree radial growth might increase by 19% in the study region, compared to the average during the period 1952–2010. Consequently, tree stem radial growth is expected to increase under a warming and wetting climatic scenario, but will decrease under drying conditions.     

芦芽山林线白杄生长季径向生长动态

高山林线作为树木分布的高度上限, 是全球范围最重要的植被过渡带之一, 其树木生长显著受到外界极端环境条件的影响。利用点状树木径向变化记录仪于2009年5-9月, 对山西省芦芽山林线组成树种白杄(Picea meyeri)生长季内树木径向生长进行了持续的动态监测。结果表明: 白杄茎干日变化主要受到树木蒸腾作用日变化的影响, 茎干呈现出白天脱水收缩与夜间吸水膨胀的循环变化; 生长季白杄径向生长可划分为3个不同的生长时段: 1)茎干水分恢复时段, 2)茎干快速生长时段, 3)茎干脱水收缩时段。在茎干水分恢复时段, 白杄茎干径向累积变化主要受到土壤含水量变化的影响。土壤温度是茎干快速生长时段影响茎干径向生长的主导环境因子, 同时它也影响着白杄茎干径向生长的开始。在茎干脱水收缩时段, 土壤温度、土壤含水量是影响茎干径向累积变化的主要环境因子。白杄径向生长最大速度出现在6月末, 其主要受到光周期(即白昼长短)影响, 是对林线处极端环境的一种适应。     

Water relations determine short time leaf growth patterns in the mangrove Avicennia marina (Forssk.) Vierh.

High‐resolution leaf growth is rarely studied despite its importance as a metric for plant performance and resource use efficiency. This is in part due to methodological challenges. Here, we present a method for in situ leaf growth measurements in a natural environment. We measured instantaneous leaf growth on a mature Avicennia marina subsp. australasica tree over several weeks. We measured leaf expansion by taking time‐lapse images and analysing them using marker tracking software. A custom‐made instrument was designed to enable long‐term field studies. We detected a distinct diel growth pattern with leaf area shrinkage in the morning and leaf expansion in the afternoon and at night. On average, the observed daily shrinkage was 37% of the net growth. Most of the net growth occurred at night. Diel leaf area shrinkage and recovery continued after growth cessation. The amount of daily growth was negatively correlated with shrinkage, and instantaneous leaf growth and shrinkage were correlated with changes in leaf turgor. We conclude that, at least in this tree species, instantaneous leaf growth patterns are very strongly linked to, and most likely driven by, leaf water relations, suggesting decoupling of short‐term growth patterns from carbon assimilation.     

Field evidences for the positive effects of aerosols on tree growth

Theoretical and eddy covariance studies demonstrate that aerosol‐loading stimulates canopy photosynthesis, but field evidence for the aerosol effect on tree growth is limited. Here, we measured in situ daily stem growth rates of aspen trees under a wide range of aerosol‐loading in China. The results showed that daily stem growth rates were positively correlated with aerosol‐loading, even at exceptionally high aerosol levels. Using structural equation modeling analysis, we showed that variations in stem growth rates can be largely attributed to two environmental variables covarying with aerosol loading: diffuse fraction of radiation and vapor pressure deficit (VPD). Furthermore, we found that these two factors influence stem growth by influencing photosynthesis from different parts of canopy. Using field observations and a mechanistic photosynthesis model, we demonstrate that photosynthetic rates of both sun and shade leaves increased under high aerosol‐loading conditions but for different reasons. For sun leaves, the photosynthetic increase was primarily attributed to the concurrent lower VPD; for shade leaves, the positive aerosol effect was tightly connected with increased diffuse light. Overall, our study provides the first field evidence of increased tree growth under high aerosol loading. We highlight the importance of understanding biophysical mechanisms of aerosol‐meteorology interactions, and incorporating the different pathways of aerosol effects into earth system models to improve the prediction of large‐scale aerosol impacts, and the associated vegetation‐mediated climate feedbacks.     

Production of dissolved organic carbon and low-molecular weight organic acids in soil solution driven by recent tree photosynthate

Dissolved organic carbon (DOC) is an important component in the terrestrial carbon cycle. Yet, the relative importance of different inputs of DOC to the soil solution remains uncertain. Here, we used a large-scale forest girdling experiment to examine how the supply of recent photosynthate to tree roots and their mycorrhizal fungi affects DOC, in particular low-molecular weight organic acids (LMWOA). We also studied effects of tree girdling on non-structural carbohydrates in microorganism, and examined the effects of freezing of soil and the presence of roots in the soil samples on soil solution DOC and LMWOA in this experiment. The concentration of DOC was reduced by 40%, while citrate was reduced by up to 90% in the soil solution by the girdling treatment. Other LMWOA such as oxalate, succinate, formate and propionate were unaffected by the girdling. We also found that girdling reduced the concentrations of trehalose (by 50%), a typical fungal sugar, and of monosaccharides (by 40%) in microorganisms in root-free soil. The effect of freezing on DOC concentrations was marked in samples from control plots, but insignificant in samples from girdled plots. Release of DOC from cell lysis after freezing was attributed equally to roots and to microorganisms. Our observations suggest a direct link from tree photosynthesis through roots and their mycorrhizal fungi to soil solution chemistry. This direct link should impact solute transport and speciation, mineral weathering and C dynamics in the soil compartment. Importantly, our finding of a substantial photosynthate driven production of DOC challenges the paradigm that DOC is mainly the result of decomposition of organic matter.     

Cerambycid girdling and water stress modify mesquite architecture and reproduction

This study evaluated the effect of an outbreak of the cerambycid beetle, Oncideres rhodosticta, on branch growth and inflorescence production of the mesquite Prosopis glandulosa var. torreyana, and on larvae mortality in girdled branches at two sites (dry and wet) in the southern Chihuahuan Desert of Mexico. We compared stem growth responses to girdling in branches of similar sized trees from both sites over 42 months. The number of larvae per girdled branch was similar between sites, indicating similar ovipositing effort regardless of water stress. However, the proportion of dead larvae was significantly lower in trees at the dry site. On average, girdling reduced 96% of the stem length at both sites. At the end of the first year, 25% of the original stubs survived at the dry site, compared to 90% at the wet site. Girdling also broke apical dominance and allowed for the development of lateral buds from the surviving stubs, which produced many fewer branches and inflorescences at the dry site compared to the wet site, where a compact crown was developed. Water stress and girdling have a combined effect on mesquite architecture and reproduction, since more stubs died at the dry site while new branches from surviving stubs developed at the wet site, recovering the original lost biomass but changing the appearance of the tree.     

Effects of bending stress on taper and growth of stems of youngEucalyptus regnans trees

A glasshouse experiment investigated the effect of bending stress on stem radial and height growth and stem taper ofEucalyptus regnans seedlings. Eighteen-week-old, potted seedlings were bent continuously for 8 weeks with a static bending stress. The bending treatment was then removed and the seedlings grown for another 12 weeks. Other seedlings were stayed vertically throughout the experiment whilst control seedlings were neither bent nor stayed. Seedlings were rotated every 2 days to prevent reaction wood developing asymmetrically in the stems of bent trees. Bent trees had higher radial growth rates, developed more tapered stems and had higher safety factors (the ratio of stem radius to the minimum radius required to prevent the tree toppling over) than unbent seedlings. They produced a band of tension wood in their stems and ceased height growth whilst bent. When bending ceased, they resumed normal radial and height growth. Unbent trees developed more cylindrical stems. There were no differences in growth behaviour between stayed and control trees. Bent and unbent trees all developed a butt swell, the taper of which was not affected by treatment. It was concluded that bending stress has substantial effects on both the size and taper of tree stems. However, the development of butt swell is independent of the bending stress applied. The results were considered in relation to biomechanical theories of tree stem development.     

Evaluation of drought response of two poplar clones (Populus x canadensis Monch 'I-214' and P. deltoides Marsh. 'Dvina') through high resolution analysis of stem growth

Different irrigation effects on stem radius variation (DeltaR) and maximum daily shrinkage (MDS) in Populus deltoides 'Dvina' and Populusxcanadensis 'I-214' were studied to assess differences in drought tolerance between clones. One-year-old trees growing in concrete tanks were submitted to two irrigation regimes (natural rainfall and irrigation) from 24 June to 10 August, and DeltaR was monitored by automatic point dendrometers. Independently of the irrigation regime, 'Dvina' showed a higher stem radial increment than 'I-214'. In both clones, the first response to changed soil water content was a significant increase in MDS, whilst DeltaR decreased about 20 d later when pre-dawn leaf water potential (Psipd) dropped below -0.4 MPa. However, they displayed different strategies to overcome drought. 'Dvina' maintained a positive DeltaR for longer than 'I-214', which had lower leaf Psipd and greater leaf abscission at the end of the drought period. After irrigation resumed, 'Dvina' showed a higher capacity to restore stem growth. 'I-214' was probably unable to recover secondary growth because of higher leaf abscission during drought stress and the production of newly expanded leaves during recovery. It is concluded that the larger radial growth of 'Dvina' derived from a better water use (carbon uptake versus water loss) than 'I-214' under limited water availability.     

河北塞罕坝樟子松径向生长动态变化及其与气象因子的关系

树木是森林生态系统的基本组成, 其生长受气象因子的影响, 基于此, 该研究通过监测樟子松(Pinus sylvestris var. mongolica)的径向生长, 研究樟子松生长日动态规律、季节动态规律及其与气象因子的关系, 探讨河北塞罕坝地区樟子松森林生态系统对气候变化的响应机制。此外, 以往研究树木生长大多数基于树轮年代学, 缺少短期树木径向生长动态的研究。该研究利用径向生长记录仪监测河北塞罕坝机械林场内樟子松连续3年(2016-2018)的树干径向动态变化。结果表明: 由于树干的水分吸收与蒸腾作用, 樟子松树干径向昼夜变化呈现季节性规律, 可划分为4个阶段: 春季萌动期、夏季生长期、秋冬交替期和冬季休眠期。塞罕坝樟子松树干径向生长开始于每年4月初; 4月初至5月中旬为水分恢复阶段; 5月中旬至7月中旬为快速生长阶段; 7月中旬至10月中旬为缓慢生长阶段; 10月中、下旬生长趋于停止, 并有树干径向收缩现象。以一天为时间尺度, 在快速生长阶段(5月初至7月中旬)樟子松径向生长主要受空气温度的影响; 缓慢生长阶段(7月中旬至10月下旬)降水量、空气温度均影响樟子松径向生长。以15天为时间尺度, 温度对樟子松径向生长的影响显著。结果显示樟子松的生长动态规律及其影响因子, 为未来樟子松生理研究提供参考时间节点, 同时在极端低温与干旱的情况下, 为半干旱地区樟子松的生长状态提供参考依据。     

Stem annual increments as ecobiological indicators in Turkey oak (Quercus cerris L.)

In recent decades, the biostatistical analysis of relationships among tree growth and macroclimatic factors has developed remarkably. In this discipline, the parametrisation of tree growth is generally based on the records of annual ring widths measured on breast height transverse sections of tree stems (ARI). The present research enables the stem volume annual increment (AVI) to be evaluated as a dendroclimatic indicator in comparison with ARI. The research is a part of a broader investigation into the ecobiological characteristics of Turkey oak in the Gargano region (southern Italy). The results indicate that AVI is no better than ARI as a means of highlighting expressive relationships among the examined macroclimatic factors (monthly precipitation and monthly averages of temperature daily maxima and minima) and the Turkey oak annual stem growth. In the examined stands, the most growthinfluencing factors are the autumn and spring rainfalls. Regarding the stem breast height radial increment, the most effective temperature factors are the June averages of daily minima and maxima.     

梨枣花果期耗水规律及其与茎直径变化的相关分析

设置4个水分处理,研究了4年生梨枣2010年及2011年花果期不同供水条件下土壤水分动态和耗水规律,分析了梨枣日耗水量与茎直径变化间的相关性,建立回归模型.结果表明:(1)2a内各处理梨枣耗水量随土壤供水量的增加而增大,其日耗水量最大值均出现在灌水后1周内;各处理果实膨大期日耗水强度大于开花坐果期.(2)2a内各处理茎直径日变化平均值(MTD)、茎直径日最大值(MXTD)均符合Logistic函数关系,MXTD与MTD在表征梨枣茎秆生长规律方面效果一致,各处理茎直径变化指标(MTD、MXTD)增长率因水分处理的不同而存在差异.(3)高水分(T1处理)条件下茎直径变化指标(MTD、MXTD、MDS(茎直径日最大收缩量)、DG(茎直径日生长量))在表征枣树耗水状况方面不敏感;在低水分(T4处理)条件下,日耗水量与茎直径日最大收缩量(MDS)相关系数较其他3个茎直径变化指标(MTD、MXTD、DG)高且达极显著水平,说明MDS能够更好的表征低水分处理的梨枣耗水规律.在此基础上建立耗水量与茎直径变化回归模型,为评价梨枣耗水状况提供依据.     

杜仲剥皮再生对生长的影响

杜仲（Ｅｕｃｏｍｍｉａ ｕｌｍｏｉｄｅｓ Ｌｌｉｖ．）剥皮后每年７月测定剥皮株和照株的径向与高度生长量,每年秋季观测这些树的落叶情况,春季观测其萌芽情况,连续观察４年。同是对剥皮３年后在原剥皮部位作第二次剥皮的植株作了同样的３年观测。结果表明,剥皮当年树的落叶提前、发芽推迟,高度和树粗的生长明显减慢,剥皮后第二年其各方面的生长状况与对照株接近,第三年剥皮株的生长与对照株相似,但剥皮株的木材年生长量     

树木年内径向生长对干旱事件的响应——以贺兰山油松为例

在贺兰山苏峪口国家森林公园,利用径向生长测量仪监测2017和2018年2个生长季内、2个海拔(2010和2330 m)油松的径向生长,研究树木径向生长对干旱的响应。结果表明: 2018年6月的干旱事件使得油松径向生长速率减慢,生长量减小;而7—8月的降水使油松的径向生长重新激活。2018年油松的径向生长主要发生在6—8月,相比2017年延长一个月。油松径向生长与气候因子的响应关系在不同海拔间没有明显的差异。生长季早期干旱对树木径向生长有抑制作用,生长季中后期降水对树木径向生长具有促进作用。该区的气候重建工作中应当充分考虑8月的气候要素对树轮宽度的影响。