Vegetation is a key component of the Earth system, absorbing approximately one-third of anthropogenic CO2 emissions1,2. Global climate change has substantially extended growing-season length (GSL) of vegetation and amplified seasonal peak photosynthesis (GPPmax) in rencent decades3–6, increasing terrestrial carbon uptake. Prevailing evidence suggests that warming alleviates climate constraints on vegetative photosynthetic activity in the northern regions7,8. However, recent studies also indicate that negative effects of warming on ecosystem carbon sequestration are emerging, casting uncertainty on the sustainability of the observed increase in terrestrial carbon uptake over the past few decades9,10. Interannual variations of gross primary productivity (GPP) are tightly related to GSL and GPPmax but GPPmax accounts for more than does GSL11,12. While a continuous upswing in peak vegetation growth remains ecologically tenable13, this positive trajectory may not be perpetual, owing to shifts in resource availability. Consequently, achieving a comprehensive understanding of the mechanisms governing peak vegetation growth assumes paramount importance in forecasting vegetative responses under forthcoming climate scenarios.
The magnitude of vegetative photosynthesis is intricately linked to the synchronization of seasonal vegetation growth with resource availability14. An optimal photosynthetic capacity is postulated to be achieved when the timing of peak photosynthetic activity (DOYGPP) aligns with the timing of seasonal peak of radiation (DOYPAR)15. However, DOYGPP may shift from DOYPAR due to constraints imposed by water availability and temperature16–18. Following the law of minimum19, DOYGPP would adjust towards those time intervals when the limiting resource, whether it be temperature or water, is more abundant.
Here, we used a recent machine-learning-derived solar-induced chlorophyll fluorescence (SIF) dataset, specifically the Reconstructed TROPOMI SIF (RTSIF)20, as a proxy for vegetation photosynthesis. The DOYRTSIF retrieved from RTSIF showed very high consistency with the DOYGPP retrieved from the GPP estimates from eddy covariance flux towers (Fig. S1). We investigated the synchrony of DOYRTSIF and DOYPAR and used their differences (ΔDOYRTSIF,PAR) as representatives of temperature and water constraints to assess climatic constraints in northern ecosystems (> 30oN)5. Our findings unveiled a notable expansion of regions constrained by limited water resources during the past two decades. We further established a general relationship predicated on mean annual temperature and precipitation, which effectively demarcated regions affected by either water or temperature constraints. Leveraging this analytical framework, we projected forthcoming alterations in water and temperature constraints and their potential repercussions on vegetation growth under various carbon emission scenarios.
Seasonal peak photosynthesis and potential climatic constraints
We first estimate the seasonal peak vegetation photosynthesis (RTSIFmax) and its corresponding timing (DOYRTSIF) using the RTSIF dataset based on a weighted spline fitting algorithm with a fixed per-pixel threshold (see Methods section). Subsequently, we assess the synchronization between DOYRTSIF and the peak radiation timing (DOYPAR) within northern ecosystems, spanning latitudes greater than 30°N, during the years 2001 to 2020, to discern the prevailing constraints imposed by temperature and water availability in these ecosystems (Methods, Extended Fig. 1). We find that the majority of northern ecosystems, comprising over 69.38% of the total, experience constraints primarily due to temperature, while the remainder are predominantly restricted by water availability (Fig. 1a). The temperature-constrained regions predominantly encompass the Arctic and cold boreal regions, while water-constrained regions are mainly located in the northern forests of North America and certain temperate regions. Similar spatial patterns of ΔDOYRTSIF,PAR are obtained using an independent dataset from the Global Ozone Monitoring Experiment-2 (GOME-2) SIF (Fig. S2). It is important to emphasize that the magnitude of seasonal peak photosynthesis is closely intertwined with the climatic constraints governing vegetation growth, wherein substantial RTSIFmax values are typically observed in close proximity to ΔDOYRTSIF,PAR equating to zero. Ecosystems operating under either temperature constraints (ΔDOYRTSIF,PAR > 0) or water constraints (ΔDOYRTSIF,PAR < 0) exhibit diminished photosynthetic capacity, reflecting limitations in resource availability (Fig. S3).
Changes in DOYRTSIF have regionally varying impacts on RTSIFmax (reflected by the opposite signs of the correlation coefficient between RTSIFmax and DOYRTSIF) which are closely related to dominant climatic constraints (Fig. 1b). Similar patterns can also be found in the partial correlations between RTSIFmax and DOYRTSIF (Fig. S4). An "early peak – larger peak" pattern occurs mainly in temperature-constrained areas because more favorable thermal conditions enable vegetation to enhance its synchronization with radiation, consequently augmenting plant growth21. Conversely, in water-constrained regions, an "early peak-smaller peak" pattern prevails. This phenomenon can be mainly attributed to summer water deficits triggered by warming-induced water constraints, which impede peak vegetation growth22,23. This is consistent with the opposite trends observed between RTSIFmax and ΔDOYRTSIF,PAR for ΔDOYRTSIF,PAR<0 and ΔDOYRTSIF,PAR>0 (Fig. S3).
Changes in water and temperature constraints in the past 20 years
A widespread shift in DOYRTSIF toward earlier in the growing season is found in 50.58% of the northern vegetated area during 2001–2020 (Fig. 2a). With a relatively stable DOYPAR (Table S2), a significant decrease of ΔDOYRTSIF,PAR is detected (-0.13 days/year, p < 0.05). However, distinct trends materialize in various climatic constrained regions: a pronounced decline is noted in temperature-constrained areas (-0.08 days/year, p < 0.1), whereas water-constrained regions exhibit no statistically significant temporal trend (Fig. S5). Over the past two decades, certain boreal ecosystems have undergone a transition from positive to negative ΔDOYRTSIF,PAR, indicating a transition from temperature-constrained to water constrained status24,25 (Fig. S6). These alterations underscore the influence of climate change, which has notably diminished the relative significance of thermal constraints in boreal vegetated regions and concurrently accentuated the role of water availability in temperate regions26–30.
SIFmax increased during the past 20 years in both temperature and water-constrained regions. However, it is noteworthy that temperature-constrained regions exhibit a steeper positive slope (Fig. 2b), with a rate of change of 4.003*10− 4 mWm− 2um− 1sr− 1/yr, compared to 1.529*10− 4 mWm− 2um− 1sr− 1/yr for water-constrained regions, as determined by a two-tailed t-test (p < 0.1). These findings align with previous observations employing different vegetation indices31–35. Given the expected ongoing warming and drying trends36,37, the continuous monitoring of ecosystems situated in water-constrained regions assumes paramount importance, as these trends could amplify water stress and, consequently, impact peak photosynthesis in these ecosystems38–40.
To appraise the capacity of contemporary terrestrial ecosystem models in capturing the ramifications of seasonal climatic constraints on vegetation's seasonal peak photosynthesis, we further conducted an assessment of the performance of 13 dynamic global vegetation models (DGVMs) that are participants in the "Trends and drivers of the regional scale sources and sinks of carbon dioxide" (TRENDY v.8)41. In congruence with empirical observations, it was found that modelled GPPmax exhibited an increasing trend in both temperature-constrained and water-constrained regions, corroborating the enhanced peak photosynthesis in northern ecosystems (Fig S7). Notably, no significant difference in the upward trends of modelled GPPmax was observed between temperature-constrained and water-constrained regions (two-tailed t-test, p = 0.79). Compared with observations, most models accurately captured the negative correlation between GPPmax and DOYGPP in temperature-constrained regions (Fig. 2c). However, four of the models (DLEM, SDGVM, CLM5.0, and ORCHIDEE) failed to reproduce the positive correlation between GPPmax and DOYGPP in water-constrained regions. This divergence in model performance implies that current terrestrial carbon cycle models may underestimate the adverse consequences of warming42.
A general relationship between precipitation and temperature in separating water and temperature limited zones
Using ΔDOYRTSIF,PAR as an indicator of the combined temperature and water constraints affecting ecosystem photosynthesis, we elucidate the interplay of mean annual precipitation (MAP) and temperature (MAT) on constraint status, as illustrated in Fig. 3a. The temperature and water constrained regions can be effectively separated using a support vector machine (SVM, see Methods section)43. Notably, the solid black line in Fig. 3a indicates the non-constrained status (ΔDOYRTSIF,PAR=0) where the relationship between mean annual temperature (oC) and precipitation (mm) is
$$\begin{array}{c}MAT=0.0138\times MAP+3.7687\#\left(1\right)\end{array}$$
The fidelity of this relationship is substantiated by calculated ΔDOYGPP,PAR and DOYPAR derived from Fluxnet site observations, further corroborating the effectiveness of the proposed MAT and MAP relationship in distinguishing water and temperature-constrained ecosystems (Fig. S8). Using this simple relationship, the predicted water- and temperature-constrained regions are almost identical to those estimated using ΔDOYRTSIF,PAR (Fig. S9). It is noteworthy that the majority of vegetation types currently persist within temperature-constrained regions, except for CSH, DBF and EBF (Fig. 3b). Over the past two decades, precipitation trends have been positive for half of the vegetation types (indicated by green arrows in Fig. 3b) and negative for the remaining half (denoted by red arrows). Notably, for vegetation types situated below the optimal line defined by Eq. (1), changes in temperature and precipitation over the last two decades appear to be converging toward the optimal line. Conversely, for CSH, DBF, and EBF, it is evident that vegetation growth is increasingly constrained by water availability.
Future changes of water and temperature constraints
Equation 1 provides a straightforward approach with robustness through time for evaluating potential alterations in the constraint status of vegetation photosynthesis in forthcoming scenarios (Table S3). To examine prospective changes in the spatial distribution of temperature and water-constrained regions by the end of the 21st century (2100), we have incorporated simulated temperature and precipitation data from 13 climate models participating in phase 6 of the Coupled Model Intercomparison Project (CMIP6) under representative scenarios SSP245 and SSP585 (Fig. 4a, 4b). Our results indicate that the reduction in temperature-constrained regions is anticipated to persist as a consequence of ongoing warming, indicating a continued amelioration in the temperature constraint on vegetation photosynthetic activity. This aligns with the conclusions drawn in several recent studies regarding the reduction of temperature-limitation on vegetation growth8,44. Conversely, the shift in water-constrained regions predominantly occurs in regions such as Central and Eastern North America, the Mediterranean, Western and Southern Asia, and East Asia. Under SSP245 and SSP585, we estimate that the water-constrained regions will expand by 4.51% and 11.13%, respectively, by 2100 (Fig. 4c).
In future scenarios, MAT is projected to rise in most terrestrial areas, contributing to an expanded occurrence of water-constrained regions. To delineate the respective influences of MAT and MAP, we have maintained either MAT or MAP at the average levels observed during the current period (2016 to 2020). This allows for the quantification of alterations in water-constrained regions attributed to variations in the other variable. As expected, the combined effects of warming and diminished precipitation play pivotal roles in the transition of regions from temperature-constrained to water-constrained status. Notably, these shifts are predominantly influenced by substantial increases in MAT, which exert a more prominent effect than changes in MAP (Fig. S10).
In summary, our study has introduced a pragmatic approach for identifying the influence of temperature and water constraints on vegetation growth, hinging on the timing of peak RTSIF and peak PAR. The analysis of ΔDOYGPP,PAR over the last two decades underscores an expansion of regions with limited water resources within northern ecosystems. Furthermore, leveraging an established relationship between MAT and MAP, capable of distinguishing between conditions of water and temperature constraints, we project the ongoing expansion of water-constrained regions throughout this century, under varying carbon emission scenarios. This evolution highlights a potential underestimation of the role of water availability constraints on vegetation photosynthesis within existing terrestrial ecosystem models, thereby underscoring the need for further research to incorporate these effects into model representations.