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There were obvious seasonal variations of monthly integrated PARt, PARd, and PARb (Fig. 4). The monthly PARt was dominated by PARd and PARb in the wet and dry seasons, respectively. There was a decrease in PARt and PARb during the wet season owing to the rainy conditions. PARd peaked in summer, with the largest value of 587.2 mol m–2. The annual kd varied from 0.57 to 0.63 during the observation period (Table 1). The maximum Ta and Tw values occurred in August, with monthly mean values of about 21.1°C and 24.0°C, respectively. The annual mean Ta varied from 14.8°C to 15.5°C (Table 1), which was cooler or near the long-term climate average (15.4°C). There was little year-to-year variation of the meteorological variables, except for precipitation. The annual precipitation fluctuated from 1210.4 to 1780.2 mm (Table 1), with more than 85% of the precipitation appearing in the wet season. The seasonal variation of VPD followed the precipitation distribution, with the largest values occurring in spring.
Figure 4. (a) Monthly integrated PARt, PARd, and PARb. (b) Monthly average Ta and Tw. (c) Monthly average VPD and monthly integrated precipitation during 2016–2020. The shaded area represents the wet season (from May to October).
Variables 2016 2017 2018 2019 2020 Precipitation (mm) 1780.2 1490.3 1494.4 1210.4 1497.3 Ta (°C) 15.5 15.1 14.8 15.2 15.3 kd 0.59 0.57 0.61 0.62 0.63 Table 1. Annual total precipitation, Ta and kd from 2016 to 2020.
The radiation is prone to changing with the variation of CI. When CI was in the range of 0–0.3, the ecosystem canopy received little radiation as a result of the high reflection of radiation by cloud particles (Fig. 5). As the sky became clearer, the increase in PARt and PARb was linearly and exponentially related to CI, respectively. As for PARd, it was also very low under overcast sky conditions (CI: 0–0.2). When CI exceeded 0.2, PARd increased with the increase in CI and peaked when CI was 0.4–0.6. Under sunny sky conditions (CI > 0.8), PARd increased owing to high PARt.
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The vegetation absorbed light quantum for photosynthesis, while the light demand might be saturated under high-level radiation. Under cloudy and sunny conditions, NEP was suppressed when PARt was greater than 1500 μmol m–2 s–1 in this study. Generally, the light saturation point was lower than that over an alpine meadow site, which was 1800 μmol m–2 s–1 (Gu et al., 2003). The response of NEP to PARt under different sky conditions varied across seasons (Fig. 6; Table 2). There were higher values of the apparent quantum yield (α) under overcast and cloudy conditions than that under sunny conditions in every season, except for winter. Compared with sunny skies, the values of α under cloudy skies increased by 366.7%, 80.8%, and 438.5% in spring, summer, and autumn, respectively; and α under overcast skies increased by 900.0%, 42.2%, and 130.8% from spring to autumn, respectively. The increase of α was higher in spring while the increase of α in summer was lower. As for Pmax, it was larger under cloudy or overcast skies in spring and autumn, and larger under sunny skies in summer and winter. Compared with sunny conditions, the increase of Pmax under cloudy conditions was 58.6%, –2.8%, 7.0%, –43.3% in the four seasons, respectively; and the increase of Pmax under overcast conditions attained 59.6%, –4.0%, 60.5%, and –62.5%, again from spring–winter, respectively.
Figure 6. Light response curves of half-hourly daytime NEP to PARt across seasons during 2016–2020. The curves were fitted using Eq. (13), and the regression coefficients (α , Pmax) are presented in Table 2.
Season Weather α (μmol μmol–1) Pmax (μmol m–2 s–1) Re, day (μmol m–2 s–1) R2 Spring Sunny 0.006 5.44 0.99 0.21 Cloudy 0.028 8.63 1.75 0.28 Overcast 0.060 8.68 2.62 0.27 Summer Sunny 0.045 16.75 4.20 0.47 Cloudy 0.081 16.28 3.63 0.48 Overcast 0.064 16.08 0.97 0.29 Autumn Sunny 0.013 11.60 0.47 0.30 Cloudy 0.070 12.41 3.23 0.33 Overcast 0.030 18.62 0.78 0.23 Winter Sunny 0.004 12.62 0.95 0.31 Cloudy 0.004 7.15 0.43 0.24 Overcast 0.002 4.73 0.11 0.07 Table 2. The light-response curve parameters under different sky conditions in four seasons during 2016–2020.
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There were obvious diurnal variations of NEP under different sky conditions (Figs. 7a–d). It was observed that NEP generally reached its maximum values around noon. The maximum value of NEP was 11.6 μmol m–2 s–1, which appeared on sunny days in summer. In some grassland ecosystems, CO2 uptake during the midday might be depressed (Wang et al., 2017). This phenomenon was not observed over the study site, mainly attributed to the different weather conditions. In the Beihai wetland, the sky conditions from spring to autumn were dominated by cloudy skies, and the solar radiation under cloudy skies was lower than that under sunny skies. The majority of sunny days throughout the year occurred in winter, while the total solar radiation received by the wetland was comparatively small during this period. Furthermore, other environmental factors also could be responsible for the depression of CO2 uptake. The stress from soil water greatly affected the diurnal pattern of CO2 flux in some grasslands (Fu et al., 2006), which did not occur in this wetland.
Figure 7. (a–d) Diurnal variations of NEP, (e–h) daily NEP for the whole day and daytime, (i–l) daily GPP for the whole day and daytime under different sky conditions across all seasons during 2016–2020.
The daily NEP under different skies across seasons was compared (Fig. 7e–h). There was strong CO2 uptake in summer and weak CO2 uptake in winter. Our results show that there was no significant difference in the daytime NEP between cloudy and sunny conditions, and a significant reduction in daytime NEP was observed under overcast conditions. Compared with sunny skies, the daily NEP for the daytime under overcast skies decreased by 26.0%, 48.5%, 13.4%, and 95.7% from spring to winter, respectively. The daily NEP for the whole day was smaller than that for the daytime as a result of the nighttime ecosystem respiration, and the daily integrated NEP might even become a carbon source under overcast skies. The daily NEP for the daytime during the wet season changed with CI, and peaked under cloudy skies (CI: 0.2–0.5) (Fig. 8). The daily GPP under different skies was also compared, and the daily GPP was larger under sunny or cloudy conditions than that under overcast conditions in four seasons (Fig. 7i–l). Compared with the daily GPP for the daytime under sunny days, the daily GPP for the daytime under overcast skies decreased by 14.8%, 35.7%, 3.1%, and 54.7% in four seasons, respectively. The daily GPP for the daytime during the wet season peaked when CI was 0.2–0.5 (Fig. 8).
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Increases in PARd, PARb, and Ta significantly promoted NEP, while an increase in VPD inhibited NEP on different timescales (Fig. 9a). The half-hourly NEP was mainly controlled by PARd, the relative contribution of PARd on NEP was 29.4% (Fig. 9b). As the timescale was extended to daily and monthly timescales, the relative contribution of PARd to NEP became 28.6% and 31.2%, respectively. The relative contribution of PARb to NEP varied across timescales, and PARb could explain 10.4%–15.0% of the variance of NEP from half-hourly to monthly timescales. The air temperature (Ta) played an important role in regulating NEP, as it could explain 18.4%–50.3% of the variance of NEP on different timescales. Around 1.9%–16.8% of the variance in NEP from half-hour to monthly timescales was determined by VPD. The wind speed (U) had little effect on NEP, explaining only 0.6%–4.9% of the variance in NEP.
Figure 9. (a) The partial correlation coefficients between NEP and environmental factors, and (b) the relative contribution of environmental factors to NEP from half-hourly to monthly timescales.
The relative importance of environmental variables to daily NEP in four seasons under different sky conditions is shown in Fig. 10. In spring, the daily NEP was mainly controlled by PARd, while PARb had a secondary impact on NEP. In summer and autumn, the STE of PARd on NEP increased with increased cloud cover. Under sunny and cloudy conditions, the STEs of PARb and Ta on NEP were greater than that of other factors. While under overcast conditions, NEP was mainly controlled by PARd. In winter, PARb was the main controlling factor of NEP under sunny and cloudy skies, and VPD played an important role in regulating NEP under overcast conditions. Overall, under sunny conditions, the STE of the PARb on NEP was the greatest, followed by Ta, VPD, and PARd. Under cloudy conditions, NEP was mainly controlled by PARd, followed by Ta, PARb, and VPD. Under overcast conditions, the STE of PARd on NEP was the largest.
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Photosynthesis was found to be considerably influenced by sky conditions (Fig. 6). The response curves of daytime NEP to PARt under different sky conditions were similar, while the response of NEP to PARt was more rapid under overcast skies than that under sunny skies. This result is in line with previous research (Oliphant et al., 2011; Li et al., 2020). There was an obvious seasonal variation in the response of photosynthesis to sky conditions, with higher α values in summer and lower α values in winter (Table 2). This variation might be related to the effect of diffuse radiation and vegetation dynamics. On the one hand, high kd in summer was beneficial to photosynthesis. On the other hand, previous research conducted over the Beihai wetland showed that the aerodynamic roughness length was larger in summer, representative of more vigorous vegetation growth during this period (Shao et al., 2022). This was conducive to enhancing α. Kanniah et al. (2013) reported the important effect of the annual grass cycle on the photosynthesis rate. They claimed that the rate of photosynthesis in the wet season was approximately double that of the dry season. Additionally, α values were found to be larger under overcast or cloudy conditions than under sunny conditions in most seasons, and the increase of α was larger in spring and smaller in summer. This might be attributed to the difference in the kd increase between spring and summer. Compared with sunny skies, kd under cloudy and overcast skies in spring increased by 188.5% and 310.4%, respectively. While kd in summer only increased by 26.3% and 53.2% under cloudy and overcast conditions, respectively.
The values of Pmax in different seasons were also compared. Overall, Pmax was the largest in summer, when was 46.9%–116.7% larger than that in other seasons. The high value of Pmax in summer might be related to sufficient vegetation and favorable environmental conditions. There were high Ta and vigorous vegetation growth in summer over the Beihai wetland, which could promote the CO2 uptake of the ecosystem. In addition, the changes in radiation composition and quality could influence the ecosystem's photosynthetic productivity (Knohl and Baldocchi, 2008). Compared with direct radiation, diffuse radiation could enhance the ecosystem CO2 uptake due to its deeper penetration, more homogeneous distribution (Oliphant et al., 2011), as well as its higher ratio of blue light (Urban et al., 2012), a phenomenon known as the diffuse fertilization effect. Therefore, a larger PARd and higher radiation quality in summer compared to other seasons contributed to a larger Pmax. Similar seasonal variations of α and Pmax could be found in other wetland sites (Otieno et al., 2012; Han et al., 2013; Zhong et al., 2016). Han et al. (2013) calculated α and Pmax from May to October over a reed wetland in the Yellow River Delta, and found both α and Pmax reached their maximum in August. Zhong et al. (2016) compared α and Pmax for all seasons over a reclaimed coastal wetland in the Yangtze Estuary, and reported that α peaked in summer and Pmax peaked in summer or autumn.
We subdivided the dataset and grouped the data into different VPD and Ta classes to test the importance of PARd. The breaking points were chosen to show comparisons between classes while keeping comparable sample sizes between classes. The Ta classes were Ta > 20°C, 15°C < Ta < 20°C, and Ta > 20°C, respectively. The VPD classes were VPD < 1 kPa and VPD > 1 kPa. Under similar Ta and VPD classes, the α and Pmax values were significantly greater under cloudy or overcast conditions compared to that under sunny skies (Table 3). This implied that PARd might be vital for NEP even under similar environmental conditions. Comparing α values under different environmental classes, a lower VPD and moderate Ta appeared to favor α enhancement. PARd had a stronger promoting effect on photosynthesis under cloudy conditions when Ta was in the range of 15°C–20°C. Zhang et al. (2020) also reported that a suitable Ta range (5°C–20°C) was favorable for improving the effects of PARd on photosynthesis.
Classes Weather α
(μmol μmol–1)Pmax
(μmol m–2 s–1)R2 VPD > 1kPa Total 0.0106 9.300 0.26 Overcast 0.0124 2.781 0.30 Cloudy 0.0192 12.84 0.36 Sunny 0.0038 2.252 0.26 VPD < 1kPa Total 0.0564 11.20 0.31 Overcast 0.0275 17.82 0.34 Cloudy 0.0456 11.64 0.38 Sunny 0.0039 5.468 0.32 Ta > 20°C Total 0.1198 13.69 0.33 Overcast 0.0754 15.98 0.46 Cloudy 0.0856 13.97 0.40 Sunny 0.0261 9.636 0.25 15°C < Ta < 20°C Total 0.1946 8.254 0.21 Overcast 0.0468 11.24 0.26 Cloudy 0.1629 9.281 0.26 Sunny 0.0079 7.170 0.30 Ta < 15°C Total 0.0079 3.564 0.25 Overcast 0.0624 5.460 0.26 Cloudy 0.0242 3.305 0.23 Sunny 0.0035 1.383 0.36 Table 3. The light-response curve parameters under different environmental conditions and different sky conditions during 2016–2020.
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Different sky conditions induce changes in radiation composition and other environmental factors, leading to a confounding effect on carbon sequestration. Previous research had showed that PARd played an important role in influencing NEP under different sky conditions (Knohl and Baldocchi, 2008; Mercado et al., 2009). In the Beihai wetland, we also found that PARd had a significant effect on NEP, and the effect of PARd on NEP varied across seasons. In spring, the plants began to grow and the vegetation canopy was low. PARb could penetrate deep vegetation canopy, and the increase of PARd has limited and insignificant impacts on NEP under cloudy and overcast conditions. As the vegetation growth became more active and the vegetation canopy became higher, the impact of PARd on NEP under cloudy and overcast increased in summer and autumn. While in winter, the promotion effect of PARd on NEP was insignificant as a result of relatively low solar radiation and withered vegetation.
Overall, the STE of PARd on NEP was larger than that of other meteorological factors, and NEP was larger under cloudy skies than those under other sky conditions. This was inconsistent with the results over some sites with low vegetation canopies. Letts et al. (2005) investigated the relationship between NEE and cloudiness over a peatland in Canada with a canopy height of 0.25 m. They found there was almost no difference in NEE in all ranges with CI higher than 0.3. The low canopy height tended toward weak light extinction and little shading of the leaves in the lower part of the canopy. The low PAR saturation levels (600–700 μmol m–2 s–1 in summer) of the dominant vegetation in the peatland was another reason for the weak impact of PARd on NEE. The moss canopy was almost always light-saturated under different skies, and enhanced PARd under cloudy sky conditions hardly increased NEE in any part of the canopy. While in the Beihai wetland, the light saturation point was larger, and the fertilization effect of PARd was very important when PARt was larger than the saturated point. Kanniah et al. (2013) reported that enhanced light use efficiency under cloudy skies was not sufficient to increase carbon sequestration due to the dramatic decline of the total radiation in tropical savannas. The decrease in canopy productivity under cloudy conditions resulting from the reduction in solar radiation could not be compensated by the enhanced canopy productivity of shaded leaves, which benefited from the increase in PARd. Alton (2008) analyzed the relationship between carbon uptake and global radiation at 38 sites of different ecosystems in FLUXNET and suggested that the carbon uptake declined dramatically when global radiation sharply decreased.
The air temperature (Ta) had an important effect on NEP, and the increase in Ta was conducive to enhancing NEP under most sky conditions. An increase in Ta not only improves enzyme activity but also enhances the electron transfer efficiency and the carboxylation rates of leaves. This was favorable for the photosynthetic rates of plants (Oliphant et al., 2011; Song et al., 2014). However, extremely high Ta might limit NEP by accelerating leaf aging, increasing leaf thickness, and leading to the closure of plant stomata (Williams et al., 2013). High Ta in summer was found to have an insignificant inhibiting effect on NEP (Fig. 10). Vapor pressure deficit (VPD) was another important factor for NEP, noting its inhibiting effect on NEP under sunny conditions over this wetland. The leaf cells of plants were strongly affected by the balance of water vapor pressure inside and outside the leaves. When VPD increased, the leaves' stomata tended to close, which resulted in a reduced intra-leaf CO2 concentration, thus limiting vegetation photosynthesis (Goodrich et al., 2015). Additionally, high VPD might restrict NEP by limiting the enzymatic processes, constraining biochemical capacity, and increasing the mesophyll resistance (Flexas et al., 2012). As the cloudiness increased, the inhibiting effects of VPD on NEP decreased, and it even had a promoting effect on NEP under overcast conditions. This might be attributed to the low VPD, typical under overcast skies, which was 0.12 ± 0.06 kPa. The ecosystem was mainly influenced by light conditions rather than water conditions under this scenario, so the limitation of NEP by VPD might be reduced. A study conducted in a tropical rainforest suggested that the ecosystem’s photosynthesis in dry years responded more significantly to an increase in VPD than that in wet years (Zhang et al., 2011).
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Changes in sky conditions had been reported to affect LUE (Oliphant et al., 2011). We found that daily LUE during the wet season reached its maximum under overcast conditions (CI: 0–0.2) (Fig. 8). Larger LUE under gloomy skies was partly caused by favorable environmental conditions. On sunny days, there were high levels of UV radiation. Plants would reduce photosynthesis and LUE by regulating pigmentation and enzyme mechanisms to protect themselves from strong radiation (Correia et al., 2005). As CI decreased, the fertilization effect of diffuse radiation could promote LUE and photosynthesis under cloudy and overcast skies (Dengel and Grace, 2010; Kanniah et al., 2013). However, a study conducted in grassland and peatland ecosystems with low leaf area index showed different results (Letts et al., 2005). It reported that LUE did not increase under cloudy skies. The inconsistency could be attributed to the difference in climate conditions (i.e., light and hydrothermal conditions) between these ecosystems.
Water use efficiency (WUE) was considered as an indicator quantifying the carbon−water coupling of terrestrial ecosystems, and it was critical for evaluating the relationship between ecosystem photosynthesis and water consumption. The daily average WUE during daytime in the wet season was 1.42
$ \pm $ 0.75 g C mm–1 H2O, which was within the range reported in other sites (0.65–5.4 g C kg–1 H2O) (Yu et al., 2008; Brümmer et al., 2012). We compared the WUE under different sky conditions, and found that WUE peaked under cloudy skies (Fig. 8). Cloudiness could reduce surface water loss by changing the amount of solar radiation and hydrothermal conditions. A previous study had shown that ET was larger under sunny skies than that under other skies (Shao et al., 2022). While GPP reached its maximum under cloudy skies due to a moderate Ta and PAR, as well as a low VPD and the associated high stomata openness of plants. The greater CO2 uptake and lower water loss under cloudy skies increased WUE. Liu et al. (2022) investigated WUE under different sky conditions over six plantation sites; they also reported that the environmental conditions under cloudy conditions were more conducive to WUE enhancement.The wetland storage of substantial organic carbon plays an important role in the global terrestrial carbon cycle and is found to be sensitive to climate change (Charman et al., 2013). Previous studies have shown that the amount of radiation incident at the ground surface, as well as other environmental variables, varied dramatically under global climate change (Wild, 2009). The climatically induced changes among these variables could greatly impact the environmental and social services of the wetlands, such as sequestering carbon and regulating runoff. Investigating the impact of sky conditions on the NEP, LUE, and WUE over the Beihai wetland, could enhance the understanding of ecosystem photosynthesis and CO2 uptake in wetlands. This not only has the potential to provide an important scientific basis for carbon modeling but also provides theoretical references for the sustainable development of wetlands and ecological environmental protection under global climate change.
Variables | 2016 | 2017 | 2018 | 2019 | 2020 |
Precipitation (mm) | 1780.2 | 1490.3 | 1494.4 | 1210.4 | 1497.3 |
Ta (°C) | 15.5 | 15.1 | 14.8 | 15.2 | 15.3 |
kd | 0.59 | 0.57 | 0.61 | 0.62 | 0.63 |