植物物候研究进展

植物物候直接反映了气候变化的影响,是植被动态模拟的关键.在遥感和模型技术的推动下,植物物候与全球变化关系的研究日益受到人们的关注.文中从植物物候与环境因子的相互关系、植物物候对全球变化的响应以及植物物候的遥感监测方面,综合论述了植物物候的研究进展,找出植被物候研究的不足,进而提出未来植被物候的研究方向.     

Seasonality of North Atlantic phytoplankton from space: impact of environmental forcing on a changing phenology (1998–2012)

Seasonal pulses of phytoplankton drive seasonal cycles of carbon fixation and particle sedimentation, and might condition recruitment success in many exploited species. Taking advantage of long‐term series of remotely sensed chlorophyll a (1998–2012), we analyzed changes in phytoplankton seasonality in the North Atlantic Ocean. Phytoplankton phenology was analyzed based on a probabilistic characterization of bloom incidence. This approach allowed us to detect changes in the prevalence of different seasonal cycles and, at the same time, to estimate bloom timing and magnitude taking into account uncertainty in bloom detection. Deviations between different sensors stressed the importance of a prolonged overlap between successive missions to ensure a correct assessment of phenological changes, as well as the advantage of semi‐analytical chlorophyll algorithms over empirical ones to reduce biases. Earlier and more intense blooms were detected in the subpolar Atlantic, while advanced blooms of less magnitude were common in the Subtropical gyre. In the temperate North Atlantic, spring blooms advanced their timing and decreased in magnitude, whereas fall blooms delayed and increased their intensity. At the same time, the prevalence of locations with a single autumn/winter bloom or with a bimodal seasonal cycle increased, in consonance with a poleward expansion of subtropical conditions. Changes in bloom timing and magnitude presented a clear signature of environmental factors, especially wind forcing, although changes on incident photosynthetically active radiation and sea surface temperature were also important depending on latitude. Trends in bloom magnitude matched changes in mean chlorophyll a during the study period, suggesting that seasonal peaks drive long‐term trends in chlorophyll a concentration. Our results link changes in North Atlantic climate with recent trends in the phenology of phytoplankton, suggesting an intensification of these impacts in the near future.     

黄浦江浮游植物群落结构及其对水环境的指示作用

2004年4月至2005年2月共6次对黄浦江的浮游植物进行了调查,共设5个采样点。每2个月在各样点进行定性、定量采集浮游植物,同时测定了水体部分理化指标。采用生物指示法、香农(Shannon-Wiener)多样性指数法和Margalef多样性指数法等对水质进行了评价。结果表明,黄浦江在夏季为V类水,其它时期为IV类;所受的有机污染较为严重,属于中度-乙型(β-中污)阶段。其指示浮游植物为颗粒直链藻(Melosira granulata)及其变种和四尾栅藻(Scenedesmus quadricauda (Turp.) Br啨b.)等。     

Floodplain vegetation phenology in the Southeast USA: Optimizing the timing of aerial imagery acquisition

Phenological patterns of wetland vegetation were studied on the Savannah River Site, Aiken, South Carolina, USA, to aid in delineation of wetland vegetation community types and plant species from aerial imagery. Relative phenology patterns were recorded for 16 dominant wetland plant species from May 1984 to June 1985 on the floodplains of three streams. These patterns were identified and compared with aerial photographs acquired in the same season as ground observations. Leafout and leaf senescence varied between and within floodplains. In spite of this variability, vegetation classes and individual species may be distinguished best during the late April leafout period.     

东北地区植被物候时序变化

植被与气候的关系非常密切,植被物候可作为气候变化的指示器。东北地区位于我国最北部,是气候变化的敏感区域,研究该区植被物候对气候变化的响应对阐明陆地生态体统碳循环具有重要意义。利用GIMMS AVHRR遥感数据集得到了东北地区阔叶林、针叶林、草原和草甸4种植被25a(1982—2006年)的物候时序变化,得出4种植被春季物候都表现出先提前后推迟的现象,秋季物候的变化则比较复杂,阔叶林和针叶林整体上呈现出秋季物候推迟的趋势,草原和草甸则表现为提前-推迟-提前的趋势。应用偏最小二乘(Partial Least Squares)回归分析了该区域植被物候与气候因子之间的关系,结果表明:春季温度与阔叶林、针叶林和草甸春季物候负相关,前一年冬季温度与草原春季物候正相关,降水与植被春季物候的关系有点复杂;4种植被秋季物候与夏季温度均呈正相关,除草原外,其余3种植被秋季物候均与夏季降水负相关。植被春季物候可能主要受温度影响,而秋季物候很可能主要受降水控制。     

迁地保育植物花期物候对迁入地气候变化的响应

以中国科学院武汉植物园内栽培的长果秤锤树(Sinojackia dolichocarpa C. J. Qi)、山白树(Sinowilsonia henryi Hemsl.)、夏腊梅(Sinocalycanthus chinensis Cheng et S. Y. Chang)、紫茎(Stewartia sinensis Rehd. et Wils.)和绒毛皂荚(Gleditsia vestita Chun et How ex B. G. Li) 5种迁地保育植物为对象,通过2008-2016年观察记录的初花期物候及整个花期长度的数据,研究花期的年际变化规律及其与迁入地武汉气候因子的相关性。结果显示:(1)从初花期来看,长果秤锤树的初花期每年提前1.25 d,紫茎的初花期每年推迟1.35 d,绒毛皂荚的初花期每年推迟1.22 d。(2)从花期长度来看,山白树的花期每年增加1.72 d,夏蜡梅的花期每年减少1.62 d,紫茎的花期每年增加0.32 d。(3)从花期与气候因子的相关性来看,年降水量、年平均相对湿度、 10℃有效积温、花前 10℃的有效积温是影响这5种植物初花期、花期长度的主要气候因子;不同物种间影响花期的主要气候因子有所差异。     

Independent effects of warming and nitrogen addition on plant phenology in the Inner Mongolian steppe

Background and Aims

Phenology is one of most sensitive traits of plants in response to regional climate warming. Better understanding of the interactive effects between warming and other environmental change factors, such as increasing atmosphere nitrogen (N) deposition, is critical for projection of future plant phenology.

Methods

A 4-year field experiment manipulating temperature and N has been conducted in a temperate steppe in northern China. Phenology, including flowering and fruiting date as well as reproductive duration, of eight plant species was monitored and calculated from 2006 to 2009.

Key Results

Across all the species and years, warming significantly advanced flowering and fruiting time by 0·64 and 0·72 d per season, respectively, which were mainly driven by the earliest species (Potentilla acaulis). Although N addition showed no impact on phenological times across the eight species, it significantly delayed flowering time of Heteropappus altaicus and fruiting time of Agropyron cristatum. The responses of flowering and fruiting times to warming or N addition are coupled, leading to no response of reproductive duration to warming or N addition for most species. Warming shortened reproductive duration of Potentilla bifurca but extended that of Allium bidentatum, whereas N addition shortened that of A. bidentatum. No interactive effect between warming and N addition was found on any phenological event. Such additive effects could be ascribed to the species-specific responses of plant phenology to warming and N addition.

Conclusions

The results suggest that the warming response of plant phenology is larger in earlier than later flowering species in temperate grassland systems. The effects of warming and N addition on plant phenology are independent of each other. These findings can help to better understand and predict the response of plant phenology to climate warming concurrent with other global change driving factors.     

Predicting a change in the order of spring phenology in temperate forests

The rise in spring temperatures over the past half‐century has led to advances in the phenology of many nontropical plants and animals. As species and populations differ in their phenological responses to temperature, an increase in temperatures has the potential to alter timing‐dependent species interactions. One species‐interaction that may be affected is the competition for light in deciduous forests, where early vernal species have a narrow window of opportunity for growth before late spring species cast shade. Here we consider the Marsham phenology time series of first leafing dates of thirteen tree species and flowering dates of one ground flora species, which spans two centuries. The exceptional length of this time series permits a rare comparison of the statistical support for parameter‐rich regression and mechanistic thermal sensitivity phenology models. While mechanistic models perform best in the majority of cases, both they and the regression models provide remarkably consistent insights into the relative sensitivity of each species to forcing and chilling effects. All species are sensitive to spring forcing, but we also find that vernal and northern European species are responsive to cold temperatures in the previous autumn. Whether this sensitivity reflects a chilling requirement or a delaying of dormancy remains to be tested. We then apply the models to projected future temperature data under a fossil fuel intensive emissions scenario and predict that while some species will advance substantially others will advance by less and may even be delayed due to a rise in autumn and winter temperatures. Considering the projected responses of all fourteen species, we anticipate a change in the order of spring events, which may lead to changes in competitive advantage for light with potential implications for the composition of temperate forests.     

Codominant water control on global interannual variability and trends in land surface phenology and greenness

Identifying the relative importance of climatic and other environmental controls on the interannual variability and trends in global land surface phenology and greenness is challenging. Firstly, quantifications of land surface phenology and greenness dynamics are impaired by differences between satellite data sets and phenology detection methods. Secondly, dynamic global vegetation models (DGVMs) that can be used to diagnose controls still reveal structural limitations and contrasting sensitivities to environmental drivers. Thus, we assessed the performance of a new developed phenology module within the LPJmL (Lund–Potsdam–Jena managed Lands) DGVM with a comprehensive ensemble of three satellite data sets of vegetation greenness and ten phenology detection methods, thereby thoroughly accounting for observational uncertainties. The improved and tested model allows us quantifying the relative importance of environmental controls on interannual variability and trends of land surface phenology and greenness at regional and global scales. We found that start of growing season interannual variability and trends are in addition to cold temperature mainly controlled by incoming radiation and water availability in temperate and boreal forests. Warming‐induced prolongations of the growing season in high latitudes are dampened by a limited availability of light. For peak greenness, interannual variability and trends are dominantly controlled by water availability and land‐use and land‐cover change (LULCC) in all regions. Stronger greening trends in boreal forests of Siberia than in North America are associated with a stronger increase in water availability from melting permafrost soils. Our findings emphasize that in addition to cold temperatures, water availability is a codominant control for start of growing season and peak greenness trends at the global scale.     

Altered geographic and temporal variability in phenology in response to climate change

Aim In response to recent climate warming, numerous studies have reported an earlier onset of spring and, to a lesser degree, a later onset of autumn, both determined from phenological observations. Here, we examine whether these reported changes have affected the synchronization of events on a regional level by examining temporal and spatial variability in phenology. In particular, we study whether years with earlier springs are associated with an altered spatial variability in phenology. Location Germany and the United Kingdom. Methods Plant phenological observations of 35 different phases (events such as flowering and leafing) collected by the German Weather Service (1951–2002) and butterfly phenological records of 29 species collected by the UK Butterfly Monitoring Scheme (1976–2003) are used. In these long‐term records, we examine the temporal (year‐to‐year) variability and the spatial (geographic or between site) variability with particular emphasis on how they vary with time of the year and with earliness or lateness of the phase. Results Early phenological events (i.e. spring) are more variable than later events, both in time and in space, although the pattern is clearer for plants than for butterflies. Confirming previous results, we find a clear relationship between the mean date of spring and summer phases and the degree to which they have become earlier. The spatial variability of spring events is greater in warmer years that have faster plant development. However, late spring and summer events do not show a consistent relationship. Autumn events are somewhat more spatially variable in years characterized by later seasons. Main conclusions This is the first examination of spatial variability of plant and animal phenological events at a multinational scale. Earlier spring events are likely to be associated with increased spatial variability in plants, although this is unlikely to also be true for summer events. If species experience differential changes in geographic variation this may disrupt interactions among them, e.g. in food webs. On the other hand, these may offer advantages for mobile species. Further research on linked species is recommended.     

Comparison of phenology trends by land cover class: a case study in the Great Basin, USA

Direct impacts of human land use and indirect impacts of anthropogenic climate change may alter land cover and associated ecosystem function, affecting ecological goods and services. Considerable work has been done to identify long‐term global trends in vegetation greenness, which is associated with primary productivity, using remote sensing. Trend analysis of satellite observations is subject to error, and ecosystem change can be confused with interannual variability. However, the relative trends of land cover classes may hold clues about differential ecosystem response to environmental forcing. Our aim was to identify phenological variability and 10‐year trends for the major land cover classes in the Great Basin. This case study involved two steps: a regional, phenology‐based land cover classification and an identification of phenological variability and 10‐year trends stratified by land cover class. The analysis used a 10‐year time series of Advanced Very High Resolution Radiometer satellite data to assess regional scale land cover variability and identify change. The phenology‐based regional classification was more detailed and accurate than national or global products. Phenological variability over the 10‐year period was high, with substantial shifts in timing of start of season of up to 9 weeks. The mean long‐term trends of montane land cover classes were significantly different from valley land cover classes due to a poor response of montane shrubland and pinyon‐juniper woodland to the early 1990s drought. The differential response during the 1990s suggests that valley ecosystems may be more resilient and montane ecosystems more susceptible to prolonged drought. This type of regional‐scale land cover analysis is necessary to characterize current patterns of land cover phenology, distinguish between anthropogenically driven land cover change and interannual variability, and identify ecosystems potentially susceptible to regional and global change.     

Spring phenology at different altitudes is becoming more uniform under global warming in Europe

Under current global warming, high‐elevation regions are expected to experience faster warming than low‐elevation regions. However, due to the lack of studies based on long‐term large‐scale data, the relationship between tree spring phenology and the elevation‐dependent warming is unclear. Using 652k records of leaf unfolding of five temperate tree species monitored during 1951–2013 in situ in Europe, we discovered a nonlinear trend in the altitudinal sensitivity (S_A, shifted days per 100 m in altitude) in spring phenology. A delayed leaf unfolding (2.7 ± 0.6 days per decade) was observed at high elevations possibly due to decreased spring forcing between 1951 and 1980. The delayed leaf unfolding at high‐elevation regions was companied by a simultaneous advancing of leaf unfolding at low elevations. These divergent trends contributed to a significant increase in the S_A (0.36 ± 0.07 days 100/m per decade) during 1951–1980. Since 1980, the S_A started to decline with a rate of ?0.32 ± 0.07 days 100/m per decade, possibly due to reduced chilling at low elevations and improved efficiency of spring forcing in advancing the leaf unfolding at high elevations, the latter being caused by increased chilling. Our results suggest that due to both different temperature changes at the different altitudes, and the different tree responses to these changes, the tree phenology has shifted at different rates leading to a more uniform phenology at different altitudes during recent decades.     

Delayed autumn phenology in the Northern Hemisphere is related to change in both climate and spring phenology

The timing of the end of the vegetation growing season (EOS) plays a key role in terrestrial ecosystem carbon and nutrient cycles. Autumn phenology is, however, still poorly understood, and previous studies generally focused on few species or were very limited in scale. In this study, we applied four methods to extract EOS dates from NDVI records between 1982 and 2011 for the Northern Hemisphere, and determined the temporal correlations between EOS and environmental factors (i.e., temperature, precipitation and insolation), as well as the correlation between spring and autumn phenology, using partial correlation analyses. Overall, we observed a trend toward later EOS in ~70% of the pixels in Northern Hemisphere, with a mean rate of 0.18 ± 0.38 days yr^?1. Warming preseason temperature was positively associated with the rate of EOS in most of our study area, except for arid/semi‐arid regions, where the precipitation sum played a dominant positive role. Interestingly, increased preseason insolation sum might also lead to a later date of EOS. In addition to the climatic effects on EOS, we found an influence of spring vegetation green‐up dates on EOS, albeit biome dependent. Our study, therefore, suggests that both environmental factors and spring phenology should be included in the modeling of EOS to improve the predictions of autumn phenology as well as our understanding of the global carbon and nutrient balances.     

Shifts in the temperature‐sensitive periods for spring phenology in European beech and pedunculate oak clones across latitudes and over recent decades

Spring phenology of temperate trees has advanced worldwide in response to global warming. However, increasing temperatures may not necessarily lead to further phenological advance, especially in the warmer latitudes because of insufficient chilling and/or shorter day length. Determining the start of the forcing phase, that is, when buds are able to respond to warmer temperatures in spring, is therefore crucial to predict how phenology will change in the future. In this study, we used 4,056 leaf‐out date observations during the period 1969–2017 for clones of European beech (Fagus sylvatica L.) and pedunculate oak (Quercus robur L.) planted in 63 sites covering a large latitudinal gradient (from Portugal ~41°N to Norway ~63°N) at the International Phenological Gardens in order to (a) evaluate how the sensitivity periods to forcing and chilling have changed with climate warming, and (b) test whether consistent patterns occur along biogeographical gradients, that is, from colder to warmer environments. Partial least squares regressions suggest that the length of the forcing period has been extended over the recent decades with climate warming in the colder latitudes but has been shortened in the warmer latitudes for both species, with a more pronounced shift for beech. We attribute the lengthening of the forcing period in the colder latitudes to earlier opportunities with temperatures that can promote bud development. In contrast, at warmer or oceanic climates, the beginning of the forcing period has been delayed, possibly due to insufficient chilling. However, in spite of a later beginning of the forcing period, spring phenology has continued to advance at these areas due to a faster satisfaction of heat requirements induced by climate warming. Overall, our results support that ongoing climate warming will have different effects on the spring phenology of forest trees across latitudes due to the interactions between chilling, forcing and photoperiod.     

Common garden comparison of the leaf‐out phenology of woody species from different native climates,combined with herbarium records,forecasts long‐term change

A well‐timed phenology is essential for plant growth and reproduction, but species‐specific phenological strategies are still poorly understood. Here, we use a common garden approach to compare biannual leaf‐out data for 495 woody species growing outdoors in Munich, 90% of them not native to that climate regime. For three species, data were augmented by herbarium dates for 140‐year‐long time series. We further meta‐analysed 107 temperate‐zone woody species in which leaf‐out cues have been studied, half of them also monitored here. Southern climate–adapted species flushed significantly later than natives, and photoperiod‐ and chilling‐ sensitive species all flushed late. The herbarium method revealed the extent of species‐specific climate tracking. Our results forecast that: (1) a northward expansion of southern species due to climate warming will increase the number of late flushers in the north, counteracting documented and expected flushing time advances; and (2) photoperiod‐ and chilling‐sensitive woody species cannot rapidly track climate warming.     

Effects of experimental shifts in flowering phenology on plant-pollinator interactions

Climate change has led to phenological shifts in flowering plants and insect pollinators, causing concern that these shifts will disrupt plant-pollinator mutualisms. We experimentally investigated how shifts in flowering onset affect pollinator visitation for 14 native perennial plant species, six of which have exhibited shifts to earlier flowering over the last 70 years and eight of which have not. We manipulated flowering onset in greenhouses and then observed pollinator visitation in the field. Five of six species with historically advanced flowering received more visits when flowering was experimentally advanced, whereas seven of eight species with historically unchanged flowering received fewer visits when flowering earlier. This pattern suggests that species unconstrained by pollinators have advanced their flowering, whereas species constrained by pollinators have not. In contrast to current concern about phenological mismatches disrupting plant-pollinator mutualisms, mismatches at the onset of flowering are not occurring for most of our study species.     

Temperature sensitivity of spring vegetation phenology correlates to within-spring warming speed over the Northern Hemisphere

The inter-annual shift of spring vegetation phenology relative to per unit change of preseason temperature, referred to as temperature sensitivity (days °C⁻¹), quantifies the response of spring phenology to temperature change. Temperature sensitivity was found to differ greatly among vegetation from different environmental conditions. Understanding the large-scale spatial pattern of temperature sensitivity and its underlying determinant will greatly improve our ability to predict spring phenology. In this study, we investigated the temperature sensitivity for natural ecosystems over the North Hemisphere (north of 30°N), based on the vegetation phenological date estimated from NDVI time-series data provided by the Advanced Very High Resolution Radiometer (AVHRR) and the corresponding climate dataset. We found a notable longitudinal change pattern with considerable increases of temperature sensitivity from inlands to most coastal areas and a less obvious latitudinal pattern with larger sensitivity in low latitude area. This general spatial variation in temperature sensitivity is most strongly associated with the within-spring warming speed (WWS; r = 0.35, p < 0.01), a variable describing the increase speed of daily mean temperature during spring within a year, compared with other factors including the mean spring temperature, spring precipitation and mean winter temperature. These findings suggest that the same magnitude of warming will less affect spring vegetation phenology in regions with higher WWS, which might partially reflect plants’ adaption to local climate that prevents plants from frost risk caused by the advance of spring phenology. WWS accounts for the spatial variation in temperature sensitivity and should be taken into account in forecasting spring phenology and in assessing carbon cycle under the projected climate warming.     

北半球中高纬度地区近30年植被春季物候温度敏感性

北半球气候变暖导致植被春季物候开始日期显著提前,温度对春季物候的促进作用是一个过程事件而非瞬时事件,且存在空间差异。该研究在以前研究的基础上,进一步分析温度对植被物候的作用方式,并探讨春季物候温度敏感性的空间特征及影响因素。利用GIMMS3g卫星植被指数产品,采用5种方法提取1982–2009年植被春季物候,并结合格网气象数据计算植被春季物候的温度敏感性,着重分析自然植被春季物候温度敏感性与环境因素的关系。结果表明,温度是北半球植被春季物候的主要制约因素,54%的像元显示温度最大效应发生在物候开始当月和之前一个月。温度主导的春季物候的像元中,91.3%的像元指示早春温度对物候开始的促进作用。植被春季物候的温度敏感性存在空间异质性,随着区域环境因素的不同,年际温度标准差、累积降水量和辐射对植被春季物候温度敏感性都具有各自或协同的调控作用。     

深层海洋浮游植物研究综述

对深层海洋(200 m)浮游生物的研究是我国海洋浮游生物研究的前沿,已有的深层海洋浮游生物知识是我国相关研究的基础。浮游植物需要阳光进行光合作用,大多数浮游植物沉降到深海中会死亡,但有些浮游植物却能在深海中生存。综述深层海洋中存活的浮游植物的研究进展,为我国的研究提供参考。在深层海洋发现的浮游植物(2"m)种类不多,共有18种,最深分布在4000 m,颗石藻的丰度大于硅藻和甲藻,最大丰度为2220个/L,出现在500 m。在1911—1985年期间,曾经发现橄榄绿细胞,由于研究手段有限,一直没有对其进行分类学研究。1985年以后的研究怀疑这些橄榄绿细胞就是聚球藻蓝细菌,聚球藻蓝细菌在太平洋、大西洋和地中海的几个站位能分布到2750 m,在深层海洋的最大丰度为3.5×105个/mL,出现在800 m。在许多海区200—700 m发现有浮游植物色素荧光的高值,但对引起高值的原因尚不清楚。浮游植物生物量在深层海洋的垂直剖面研究不多,已有的研究表明浮游植物生物量在300 m以深为0.001—0.1"g C/L,且其随深度增加而降低。     

Satellite observed reversal in trends of spring phenology in the middle-high latitudes of the Northern Hemisphere during the global warming hiatus

The start of the growing season (SOS) is essential to track the responses of vegetation to climate change. However, recent findings on whether the SOS in the middle-high latitudes of the Northern Hemisphere (NH) continued to advance or reversed during the global warming hiatus were not consistent. It is necessary to investigate the causes of this controversy and to examine the relationship between the SOS and preseason temperature trends. To this end, we first applied four widely used phenology extraction methods to derive the SOS from the GIMMS NDVI3g dataset and then used the ensemble empirical modal decomposition (EEMD) method to extract the nonlinear trends of the SOS and preseason temperature. Our results clarify, for the first time, that the limitations of the linear assumption-based trend analysis methods are an important but overlooked cause of the discrepancies among existing studies on whether the SOS was advanced or delayed in the NH (>30° N) during the global warming hiatus. We further revealed the range of the mismatches between the SOS and preseason temperature trends at the latitude, altitude and biome levels. Specifically, we discovered that the SOS in the NH (>30° N) obtained by the four phenology extraction methods showed a significant reversal from advance to delay during the global warming hiatus, and the corresponding average rate of change was very small. The area showing increasing preseason temperatures decreased during the global warming hiatus, but it always occupied most of the NH (>30° N). However, delayed SOS trends were dominant in the NH from 50° N to 60° N, above 3000 m and in biomes other than TBMF and BF. Accordingly, using an EEMD-like approach to evaluate the changes in the SOS and preseason temperature is necessary for improving our understanding of the changes in the SOS and their association with climate.