Reduced early growing season freezing resistance in alpine treeline plants under elevated atmospheric CO2

The frequency of freezing events during the early growing season and the vulnerability to freezing of plants in European high‐altitude environments could increase under future atmospheric and climate change. We tested early growing season freezing sensitivity in 10 species, from four plant functional types (PFTs) spanning three plant growth forms (PGFs), from a long‐term in situ CO₂ enrichment (566 vs. 370 ppm) and 2‐year soil warming (+4 K) experiment at treeline in the Swiss Alps (Stillberg, Davos). By additionally tracking plant phenology, we distinguished indirect phenology‐driven CO₂ and warming effects from direct physiology‐related effects on freezing sensitivity. The freezing damage threshold (lethal temperature 50) under ambient conditions of the 10 treeline species spanned from ?6.7±0.3 °C (Larix decidua) to ?9.9±0.6 °C (Vaccinium gaultherioides). PFT, but not PGF, explained a significant amount of this interspecific variation. Long‐term exposure to elevated CO₂ led to greater freezing sensitivity in multiple species but did not influence phenology, implying that physiological changes caused by CO₂ enrichment were responsible for the effect. The elevated CO₂ effect on freezing resistance was significant in leaves of Larix, Vaccinium myrtillus, and Gentiana punctata and marginally significant in leaves of Homogyne alpina and Avenella flexuosa. No significant CO₂ effect was found in new shoots of Empetrum hermaphroditum or in leaves of Pinus uncinata, Leontodon helveticus, Melampyrum pratense, and V. gaultherioides. Soil warming led to advanced leaf expansion and reduced freezing resistance in V. myrtillus only, whereas Avenella showed greater freezing resistance when exposed to warming. No effect of soil warming was found in any of the other species. Effects of elevated CO₂ and soil warming on freezing sensitivity were not consistent within PFTs or PGFs, suggesting that any future shifts in plant community composition due to increased damage from freezing events will likely occur at the individual species level.     

Effects of seasonal snow on the growing season of temperate vegetation in China

Variations in seasonal snowfall regulate regional and global climatic systems and vegetation growth by changing energy budgets of the lower atmosphere and land surface. We investigated the effects of snow on the start of growing season (SGS) of temperate vegetation in China. Across the entire temperate region in China, the winter snow depth increased at a rate of 0.15 cm yr^?1 (P = 0.07) during the period 1982–1998, and decreased at a rate of 0.36 cm yr^?1 (P = 0.09) during the period 1998–2005. Correspondingly, the SGS advanced at a rate of 0.68 day yr^?1 (P < 0.01) during 1982–1998, and delayed at a rate of 2.13 day yr^?1 (P = 0.07) during 1998–2005, against a warming trend throughout the entire study period of 1982–2005. Spring air temperature strongly regulated the SGS of both deciduous broad‐leaf and coniferous forests, whereas the winter snow had a greater impact on the SGS of grassland and shrubs. Snow depth variation combined with air temperature contributed to the variability in the SGS of grassland and shrubs, as snow acted as an insulator and modulated the underground thermal conditions. In addition, differences were seen between the impacts of winter snow depth and spring snow depth on the SGS; as snow depths increased, the effect associated went from delaying SGS to advancing SGS. The observed thresholds for these effects were snow depths of 6.8 cm (winter) and 4.0 cm (spring). The results of this study suggest that the response of the vegetation's SGS to seasonal snow change may be attributed to the coupling effects of air temperature and snow depth associated with the underground thermal conditions.     

No increase in alpine snowbed productivity in response to experimental lengthening of the growing season

Climate change effects on snow cover and thermic regime in alpine tundra might lead to a longer growing season, but could also increase risks to plants from spring frost events. Alpine snowbeds, i.e. alpine tundra from late snowmelt sites, might be particularly susceptible to such climatic changes. Snowbed communities were grown in large monoliths for two consecutive years, under different manipulated snow cover treatments, to test for effects of early (E) and late (L) snowmelt on dominant species growth, plant functional traits, leaf area index (LAI) and aboveground productivity. Spring snow cover was reduced to assess the sensitivity of snowbed alpine species to severe early frost events, and dominant species freezing temperatures were measured. Aboveground biomass, productivity, LAI and dominant species growth did not increase significantly in E compared to L treatments, indicating inability to respond to an extended growing season. Edapho‐climatic conditions could not account for these results, suggesting that developmental constraints are important in controlling snowbed plant growth. Impaired productivity was only detected when harsher and more frequent frost events were experimentally induced by early snowmelt. These conditions exposed plants to spring frosts, reaching temperatures consistent with the estimated freezing points of the dominant species (～?10 °C). We conclude that weak plasticity in phenological response and potential detrimental effects of early frosts explain why alpine tundra from snowbeds is not expected to benefit from increased growing season length.     

Long‐term enhanced winter soil frost alters growing season CO2 fluxes through its impact on vegetation development in a boreal peatland

At high latitudes, winter climate change alters snow cover and, consequently, may cause a sustained change in soil frost dynamics. Altered winter soil conditions could influence the ecosystem exchange of carbon dioxide (CO₂) and, in turn, provide feedbacks to ongoing climate change. To investigate the mechanisms that modify the peatland CO₂ exchange in response to altered winter soil frost, we conducted a snow exclusion experiment to enhance winter soil frost and to evaluate its short‐term (1–3 years) and long‐term (11 years) effects on CO₂ fluxes during subsequent growing seasons in a boreal peatland. In the first 3 years after initiating the treatment, no significant effects were observed on either gross primary production (GPP) or ecosystem respiration (ER). However, after 11 years, the temperature sensitivity of ER was reduced in the treatment plots relative to the control, resulting in an overall lower ER in the former. Furthermore, early growing season GPP was also lower in the treatment plots than in the controls during periods with photosynthetic photon flux density (PPFD) ≥800 μmol m^?2 s^?1, corresponding to lower sedge leaf biomass in the treatment plots during the same period. During the peak growing season, a higher GPP was observed in the treatment plots under the low light condition (i.e. PPFD 400 μmol m^?2 s^?1) compared to the control. As Sphagnum moss maximizes photosynthesis at low light levels, this GPP difference between the plots may have been due to greater moss photosynthesis, as indicated by greater moss biomass production, in the treatment plots relative to the controls. Our study highlights the different responses to enhanced winter soil frost among plant functional types which regulate CO₂ fluxes, suggesting that winter climate change could considerably alter the growing season CO₂ exchange in boreal peatlands through its effect on vegetation development.     

A simulation of the importance of length of growing season and canopy functional properties on the seasonal gross primary production of temperate alpine meadows

Background and Aims: Along snowmelt gradients, the canopies of temperate alpine meadowsdiffer strongly in their structural and biochemical properties.Here, a study is made of the effects of these canopy dissimilaritiescombined with the snow-induced changes in length of growingseason on seasonal gross primary production (GPP). Methods: Leaf area index (LAI) and community-aggregated values of leafangle and leaf nitrogen content were estimated for seven alpineplant canopies distributed along a marked snowmelt gradient,and these were used as input variables in a sun–shadecanopy bulk-photosynthesis model. The model was validated forplant communities of early and late snowmelt sites by measuringthe instantaneous CO₂ fluxes with a canopy closed-chamber technique.A sensitivity analysis was conducted to estimate the relativeimpact of canopy properties and environmental factors on thedaily and seasonal GPP. Key Results: Carbon uptake was primarily related to the LAI and total canopynitrogen content, but not to the leaf angle. For a given levelof photosynthetically active radiation, CO₂ assimilation washigher under overcast conditions. Sensitivity analysis revealedthat increase of the length of the growing season had a highereffect on the seasonal GPP than a similar increase of any otherfactor. It was also found that the observed greater nitrogencontent and larger LAI of canopies in late-snowmelt sites largelycompensated for the negative impact of the reduced growing season. Conclusions: The results emphasize the primary importance of snow-inducedchanges in length of growing season on carbon uptake in alpinetemperate meadows. It was also demonstrated how using leaf-traitvalues of the dominants is a useful approach for modelling ecosystemcarbon-cycle-related processes, particularly when continuousmeasurements of CO₂ fluxes are technically difficult. The studythus represents an important step in addressing the challengeof using a plant functional-trait approach for biogeochemicalmodelling.     

High tolerance of repeated heatwaves in Australian native plants

Heatwaves, with increases in day and night time temperatures, are predicted to increase in frequency. We investigated the response of forbs, shrubs, grasses and non‐grass monocotyledons from warm temperate environments in Australia to repeated heatwaves to determine if responses differed with growth form and whether the addition of hot night temperatures influenced the ability of species to grow and acclimate. Plants were subjected to 3, 3‐day heatwaves comprising either hot days and nights or hot days and cool nights, with control plants maintained under cool days and nights. All species were thermotolerant to repeated heatwaves, although two species showed lower biomass under heat treatments, indicating repeated heatwaves influenced function in some, but not all species. While there was evidence of photosynthetic damage in some species, these recovered by the end of the experiment. While grasses and one herb showed some evidence of photosystem acclimation, increases in the threshold temperatures for membrane breakdown only occurred in one shrub. Leaf sacrifice in grasses was significantly increased after repeated heatwaves suggesting that fuel loads in grass communities will increase in the future. These results indicate high resilience for these Australian native warm temperate plants although lower growth rates in some species after heatwaves may result in changes to community composition.     

Low-temperature resistance in Polylepis tarapacana, a tree growing at the highest altitudes in the world

The Polylepis tarapacana forests found in Bolivia are unique with respect to their altitudinal distribution (4200–5200 m). Given the extreme environmental conditions that characterize these altitudes, this species has to rely on distinct mechanisms to survive stressful temperatures. The purpose of this study was to determine low‐temperature resistance mechanisms in P. tarapacana. Tissue was sampled for carbohydrate and proline contents and micro‐climatic measurements were made at two altitudes, 4300 and 4850 m, during both the dry cold and wet warm seasons. Supercooling capacity (?3 to ?6 °C for the cold dry and ?7 to ?9 °C for the wet warm season) and injury temperatures (?18 to ?23 °C for both seasons), determined in the laboratory, indicate that P. tarapacana is a frost‐tolerant species. On the other hand, an increase in supercooling capacity, as the result of significant increase in total soluble sugar and proline contents, occurs during the wet warm season as a consequence of higher metabolic activity. Hence, P. tarapacana, a frost‐tolerant species during the colder unfavourable season, is able to avoid freezing during the more favourable season when minimum night‐time temperatures are not as extreme.     

Leaf rosette closure in the alpine rock species Saxifraga paniculata Mill.: significance for survival of drought and heat under high irradiation

The significance of leaf rosette closure for survival of drought and heat under high irradiation on alpine rock sites was investigated in the cushion forming rosette plant, Saxifraga paniculata Mill. With decreasing water content the leaves fold over the rosette centre reducing reversibly the evaporative leaf surface area by 80%. Internal water redistribution driven by an osmotic gradient from older to younger leaves occurs. The oldest leaves dry out to promote the survival of the individual. Leaf temperatures above 45 °C (which match heat tolerance limits 45–57 °C; LT₅₀) co-occurred with low substrate water potentials (less than – 0·5 MPa) on 11·3% of summer days. Shading by leaves can be crucial to surviving high temperatures as it keeps the rosette centre up to 10 °C colder. Mutual shading prevented sustained drought-induced photoinhibition in upper leaf surfaces at relative water contents below 60%. In exposed lower leaf surfaces restoration of photosystem II took several days. Leaf temperatures above 40 °C (21·3% of summer days) induced photoinhibition in situ. Periods with sufficient water supply can be fully utilized as rehydration is fast ( < 12 h) and exposes the upper leaf surfaces that showed only minor photoinhibition. By reversible leaf rosette closure environmental extremes that otherwise could exceed tolerance are efficiently avoided.     

Spring phenology rather than climate dominates the trends in peak of growing season in the Northern Hemisphere

Shifts in plant phenology regulate ecosystem structure and function, which feeds back to the climate system. However, drivers for the peak of growing season (POS) in seasonal dynamics of terrestrial ecosystems remain unclear. Here, spatial–temporal patterns of POS dynamics were analyzed by solar-induced chlorophyll fluorescence (SIF) and vegetation index in the Northern Hemisphere over the past two decades from 2001 to 2020. Overall, a slow advanced POS was observed in the Northern Hemisphere, while a delayed POS distributed mainly in northeastern North America. Trends of POS were driven by the start of growing season (SOS) rather than pre-POS climate both at hemisphere and biome scale. The effect of SOS on the trends in POS was the strongest in shrublands while the weakest in evergreen broad-leaved forest. These findings highlight the crucial role of biological rhythms rather than climatic factors in exploring seasonal carbon dynamics and global carbon balance.     

Temperature stress and survival ability of Mediterranean sclerophyllous plants

ABSTRACT

The Mediterranean climate with hot and dry summer periods, and low winter temperatures and episodic frosts in northern, altitudinal and continental districts, demands from evergreen broadleaved woody plants an adequate and flexible acclimation to the climatic constraints.

In this brief survey on some responses of Mediterranean sclerophylls to temperature stress, the following is presented and discussed: criteria for cold and heat limits of photosynthetic function; winter depression and summer photoinactivation of photosynthesis; peculiar patterns of tissue freezing of scleromorphous leaves and limits of frost resistance of various plant parts and ontogenetic stages; heat impairment of chloroplasts and thermotolerance of sclerophyllous species; survival capacity and recovery after damage. Risks of damage to plants in relation to stressful temperatures in Mediterranean regions are estimated. Cold stress and drought stress indices, according to Mitrakos (1980), have been applied to characterise different localities in Italy. Additionally, a heat stress index for the Mediterranean region is proposed. Future research topics are suggested.     

Effects of frost hardening, dehardening and freezing stress on in vivo chlorophyll fluorescence of seedlings of Scots pine (Pinus sylvestris L.)

Abstract. The kinetics of in vivo chlorophyll fluorescence of photosystem II (PS II) was measured at room temperature and 77 K during frost hardening of seedlings of Scots pine (Pinus sylvestris L.), and after exposure of frost-hardened shoots to sub-freezing temperatures. A more pronounced decrease in variable fluorescence yield for the upper exposed than for the lower shaded surface of the needles suggested that some photoinhibition occurred during prolonged frost hardening at 50 μmol photons m^?2 s^?1 and 4°C. Reversible inhibition of photosynthesis after exposure to sub-freezing temperatures was initially manifested as an increase of steady-state energy-dependent fluorescence quenching (q_E) and a reduction in the rate of O₂ evolution. Further inhibition after treatment at still lower temperatures caused a progressive decline of steady-state photochemical quenching (q_Q) and the rate of O₂ evolution, whereas q_E remained high. This implies an inactivation of enzymes in the photosynthetic carbon reduction cycle decreasing the consumption of ATP and NADPH, which is likely to cause an increase of membrane energization and a reduction of the primary electron acceptor (Q_A) of PS II. Alternatively, the changes in q_Q and q_E might be attributed to an inhibition of photophosphorylation. Severe, irreversible damage to photosynthesis resulted in a suppression of q_E and of variable fluorescence yield, probably because the photochemical efficiency of PS II was impaired. Changes in the fast fluorescence kinetics at room temperature after severe freezing damage were interpreted as an inhibition of the electron flow from Q_A to the plastoquinone pool. It is suggested that irreversible freezing injury to needles of frost-hardened P. sylvestris causes damage to the Q_B,-protein.     

Application of heat stress in situ demonstrates a protective role of irradiation on photosynthetic performance in alpine plants

The impact of sublethal heat on photosynthetic performance, photosynthetic pigments and free radical scavenging activity was examined in three high mountain species, Rhododendron ferrugineum, Senecio incanus and Ranunculus glacialis using controlled in situ applications of heat stress, both in darkness and under natural solar irradiation. Heat treatments applied in the dark reversibly reduced photosynthetic performance and the maximum quantum efficiency of photosystem II (F_v/F_m), which remained impeded for several days when plants were exposed to natural light conditions subsequently to the heat treatment. In contrast, plants exposed to heat stress under natural irradiation were able to tolerate and recover from heat stress more readily. The critical temperature threshold for chlorophyll fluorescence was higher under illumination (T_c^′) than in the dark (T_c). Heat stress caused a significant de‐epoxidation of the xanthophyll cycle pigments both in the light and in the dark conditions. Total free radical scavenging activity was highest when heat stress was applied in the dark. This study demonstrates that, in the European Alps, heat waves can temporarily have a negative impact on photosynthesis and, importantly, that results obtained from experiments performed in darkness and/or on detached plant material may not reliably predict the impact of heat stress under field conditions.     

川西高山林线三种灌木凋落叶分解中的无脊椎动物多样性

以无脊椎动物为主体的土壤动物是影响凋落物分解的重要生物因素,对维持陆地生态系统物质循环和能量流动具有重要作用。高山林线交错带是高山植被垂直带谱中重要的过渡区域,拥有比相邻生态系统更高的生境复杂性和物种多样性。林线上温度波动和冻融循环频率显著高于针叶林,为了了解林线交错带上环境差异对凋落物分解过程中的土壤动物群落结构和多样性的影响,采用凋落物分解袋的方法,于高山生态系统的两个主要时期,即雪被末期和生长季末期,研究了林线主要代表性灌木——高山柳(Salix cupularis)、高山杜鹃(Rhododendron lapponicum)和红毛花楸(Sorbus rufopilosa)凋落叶分解的土壤动物多样性特征。结果表明:凋落物中的无脊椎动物群落多样性及个体、类群密度随物种、海拔梯度和季节而变化,且季节差异对无脊椎动物多样性的影响比物种和海拔梯度更显著。3个因子的交互作用不仅影响土壤动物群落多样性和均匀度,而且影响群落个体密度和类群密度。雪被末期,凋落物中的无脊椎动物多样性指数H、均匀度指数J及丰富度指数D以针叶林最高,优势度指数C以林线最高;生长季节末期的无脊椎动物类群密度和个体密度显著高于雪被末期。总体上,凋落物中的无脊椎动物群落丰富度以生长季末期最高,林线较针叶林丰富。这意味着,未来气候变暖情景下,灌丛密度增加,凋落物输入量增大,可能导致无脊椎动物多样性增加。     

CO2 enrichment predisposes foliage of a eucalypt to freezing injury and reduces spring growth

Seedlings of Eucalyptus pauciflora, were grown in open-top chambers fumigated with ambient and elevated [CO₂], and were divided into two populations using 10% light transmittance screens. The aim was to separate the effects of timing of light interception, temperature and [CO₂] on plant growth. The orientation of the screens exposed plants to a similar total irradiance, but incident during either cold mornings (east-facing) or warm afternoons (west-facing). Following the first autumn freezing event elevated CO₂-grown plants had 10 times more necrotic leaf area than ambient CO₂ plants. West-facing plants had significantly greater (25% more) leaf damage and lower photochemical efficiency (F_v/F_m) in comparison with east-facing plants. Following a late spring freezing event east-facing elevated CO₂ plants suffered a greater sustained loss in F_v/F_m than west-facing elevated CO₂- and ambient CO₂-grown plants. Stomatal conductance was lower under elevated CO₂ than ambient CO₂ except during late spring, with the highest leaf temperatures occurring in west-facing plants under elevated CO₂. These higher leaf temperatures apparently interfered with cold acclimation thereby enhancing frost damage and reducing the ability to take advantage of optimal growing conditions under elevated CO₂.     

不同类型喀斯特植物的荧光特征及抗旱性比较

利用叶绿素荧光技术观测了五个不同类型的喀斯特植物翅荚香槐、大盔凤仙、红背叶、牛耳朵和青冈栎在不同作用光强下的光能利用特征,并且对这五种类型植物在PEG诱导水分胁迫下的抗旱性作了比较。结果表明,随着作用光强的增加,这五种植物的光化学反应能力(qP)逐渐降低,非光化学耗散作用(NPQ)明显增加,同时PSⅡ有效光化学效率(Fv′/Fm′)随之下降,导致PSⅡ电子传递量子效率(ΦPSⅡ)也明显下降,但在相同作用光强下这五个类型植物的荧光特征无明显差异。在PEG诱导水分胁迫的实验中,牛耳朵、青冈栎在PEG处理后Fv/Fm变化不明显,而红背叶Fv/Fm值下降程度最大,其次为翅荚香槐、大盔凤仙。由此推测,牛耳朵和青冈栎的耐干旱能力最强,红背叶抗旱能力最弱,翅荚香槐、大盔凤仙居中。     

灌丛移除对荒漠齿肋赤藓越冬过程中生理生化特性的影响

灌丛和生物土壤结皮镶嵌分布是古尔班通古特沙漠一种重要的地表覆被类型, 灌丛的存在不仅为小型动物提供了庇护场所, 也为隐花植物及部分草本植物创造了良好的生存条件。然而在初冬时期移除灌丛是否会影响这些隐花植物的越冬及生长还不得而知。该研究通过模拟放牧及鼠害, 移除50%双穗麻黄(Ephedra distachya)灌丛、移除全部双穗麻黄灌丛及自然对照, 测定齿肋赤藓(Syntrichia caninervis)植株脯氨酸、可溶性糖和可溶性蛋白含量, 以及丙二醛(MDA)含量和超氧化物歧化酶(SOD)、过氧化物酶(POD)和过氧化氢酶(CAT) 3种抗氧化酶活性, 并测定原位光化学效率, 探讨初冬灌丛丧失对荒漠藓类植物越冬的影响。结果表明, 在积雪期3种处理下齿肋赤藓的游离脯氨酸和可溶性糖含量之间差异并不显著, 但去灌丛后其MDA含量、POD、SOD和CAT 3种抗氧化酶活性均显著高于自然灌丛下齿肋赤藓的相关生理指标。即使是在融雪期, 去灌丛下齿肋赤藓的可溶性糖含量及POD和CAT活性仍显著高于自然灌丛, 而可溶性蛋白明显较低, 这可能是由于灌丛的丧失造成的温度波动加剧了冬季低温对藓类植物的伤害。灌丛的部分移除(50%)对齿肋赤藓的生理生化特性影响不显著, 就积雪融化期叶绿素荧光活性持续时间而言, 与自然灌丛和移除50%灌丛相比, 完全移除灌丛的齿肋赤藓植株叶绿素荧光活性持续时间显著缩短。这可能是由于灌丛移除导致其UV-B辐射增加及“湿岛效应”消失所致, UV-B辐射的增加加剧了对植物的伤害, 而春季融雪期保水能力的下降也是其叶绿素荧光活性时间缩短的重要原因。     

The relation between membrane lipid phase separation and frost tolerance of cereals and other cool climate plant species

Abstract. The temperatures at which liposomes prepared from membrane phospholipids begin to phase separate were compared to the temperatures at which intact plants were damaged. Woody perennials tolerated temperatures below which their membrane phospholipids began to phase separate. By contrast, rye and wheat seedlings were damaged about 25°C above their phase separation temperature. Differences in tolerance among cultivars pre-hardened to frost were reflected by changes of the phase separation temperature. The results support the notion that alterations in membrane lipid composition are associated with frost hardening. A correlation between the temperature of phase separation and frost tolerance suggests that lipid properties may influence freezing tolerance of cereals; however, the lethal event is apparently not phase separation of the membrane phospholipids.     

Importance of recent shifts in soil thermal dynamics on growing season length, productivity, and carbon sequestration in terrestrial high-latitude ecosystems

In terrestrial high‐latitude regions, observations indicate recent changes in snow cover, permafrost, and soil freeze–thaw transitions due to climate change. These modifications may result in temporal shifts in the growing season and the associated rates of terrestrial productivity. Changes in productivity will influence the ability of these ecosystems to sequester atmospheric CO₂. We use the terrestrial ecosystem model (TEM), which simulates the soil thermal regime, in addition to terrestrial carbon (C), nitrogen and water dynamics, to explore these issues over the years 1960–2100 in extratropical regions (30–90°N). Our model simulations show decreases in snow cover and permafrost stability from 1960 to 2100. Decreases in snow cover agree well with National Oceanic and Atmospheric Administration satellite observations collected between the years 1972 and 2000, with Pearson rank correlation coefficients between 0.58 and 0.65. Model analyses also indicate a trend towards an earlier thaw date of frozen soils and the onset of the growing season in the spring by approximately 2–4 days from 1988 to 2000. Between 1988 and 2000, satellite records yield a slightly stronger trend in thaw and the onset of the growing season, averaging between 5 and 8 days earlier. In both, the TEM simulations and satellite records, trends in day of freeze in the autumn are weaker, such that overall increases in growing season length are due primarily to earlier thaw. Although regions with the longest snow cover duration displayed the greatest increase in growing season length, these regions maintained smaller increases in productivity and heterotrophic respiration than those regions with shorter duration of snow cover and less of an increase in growing season length. Concurrent with increases in growing season length, we found a reduction in soil C and increases in vegetation C, with greatest losses of soil C occurring in those areas with more vegetation, but simulations also suggest that this trend could reverse in the future. Our results reveal noteworthy changes in snow, permafrost, growing season length, productivity, and net C uptake, indicating that prediction of terrestrial C dynamics from one decade to the next will require that large‐scale models adequately take into account the corresponding changes in soil thermal regimes.