Stirling C M, Heddell-Cowie M, Jones M L, Ashenden T W, Sparks T H
Institute of Terrestrial Ecology, Bangor, Gwynedd LL57 2UP, UK.
To whom correspondence should be addressed.
New Phytol. 1998 Oct;140(2):343-354. doi: 10.1046/j.1469-8137.1998.00273.x.
Whereas much is known of the short-term growth response to elevated atmospheric CO concentrations, [CO ] , there is relatively little information on how the response of native species is modified by temperature, despite the fact that an increase in global mean temperature is expected to accompany the rise in [CO ]. In this study, five functionally related annual native species were exposed to different combinations of ambient and elevated [CO ] and temperatures in order to assess their response in terms of growth and allometry. Fast-growing annuals were selected for the study because their growth responses could be assessed over a major portion of the plant's life cycle and in as short a period as 8 wk. Plants were grown in eight hemi-spherical glasshouses, programmed to track outside ambient conditions and provide a replicated experimental design. Treatments comprised (i) current ambient [CO ] and temperature, (ii) elevated [CO ] (ambient+34 kPa), and ambient temperature (iii) ambient [CO ] and elevated temperature (ambient+3°C) and (iv) elevated [CO ] and elevated temperature (T°C ). All five species responded positively to [CO ] , although the response was statistically significant for only one, Poa annua L. Averaged over all five species, [CO ] increased total plant biomass by 25% (P=0·005) at 56 d, reflecting a proportionally greater increase in leaf and stem mass relative to root weight. Elevated [CO ] had no effect on leaf area, either at the individual species level or overall. Elevated T°C, by contrast, had little effect on shoot growth but increased root mass on average by 43% and leaf area by 22%. Few interactions between elevated [CO ] and T°C were observed, with the CO response generally greater at elevated than ambient T°C. Both [CO ] and T°C resulted in a transient increase in relative growth rate, (rgr), during the first 14 d exposure and a 3°C increase in temperature had no effect on the duration of the response. CO stimulation of growth operated through a sustained increase in net assimilation rate. (nar), although the potential benefit to rgr was offset by a concurrent decline in leaf area ratio (lar), as a result of a decrease in leaf area per unit leaf mass (sla). The response to T°C was generally opposite of that to [CO ] . For example, T°C increased lar through an increase in sla and this, rather than any effect on nar, was the major factor responsible for the stimulation of rgr. Allometric analysis of CO effects revealed that changes in allocation observed at individual harvests were due solely to changes associated with plant size. Elevated T°C, by contrast, had a direct effect on allocation patterns to leaves, with an increase in leaf area expansion relative to whole plant mass during the initial stages of growth and subsequent increased allocation of biomass away from leaves to other regions of the plant. No change in the allometric relation between roots and shoots were observed at either elevated [CO ] or T°C. We conclude, therefore, that allocation of biomass and morphological characteristics such as sla, are relatively insensitive to [CO ], at least when analysed at the whole-plant level, and where changes have been observed, these are the product of comparing plants of the same age but different size.
尽管我们对大气中二氧化碳浓度升高([CO₂])下的短期生长响应了解颇多,但关于温度如何改变本地物种的响应,相关信息却相对较少,尽管预计全球平均温度的升高会伴随[CO₂]的上升。在本研究中,选取了五个功能相关的一年生本地物种,使其暴露于环境和升高的[CO₂]及温度的不同组合下,以评估它们在生长和异速生长方面的响应。选择快速生长的一年生植物进行研究,是因为它们的生长响应可以在植物生命周期的大部分时间内进行评估,且最短只需8周。植物种植在八个半球形温室中,这些温室被设置为跟踪室外环境条件并提供重复的实验设计。处理包括:(i)当前环境[CO₂]和温度;(ii)升高的[CO₂](环境浓度 + 34 kPa)和环境温度;(iii)环境[CO₂]和升高的温度(环境温度 + 3°C);以及(iv)升高的[CO₂]和升高的温度(T°C)。所有五个物种对[CO₂]均有正向响应,尽管只有一年生早熟禾(Poa annua L.)的响应具有统计学显著性。在所有五个物种中进行平均,[CO₂]在56天时使总植物生物量增加了25%(P = 0·005),这反映出相对于根重,叶和茎质量的增加比例更大。升高的[CO₂]无论是在单个物种水平还是总体水平上,对叶面积均无影响。相比之下,升高的温度(T°C)对地上部生长影响较小,但平均使根质量增加了43%,叶面积增加了22%。在升高的[CO₂]和T°C之间未观察到明显的相互作用,一般而言,在升高的温度下[CO₂]的响应比在环境温度下更大。[CO₂]和T°C在最初暴露的14天内均导致相对生长速率(rgr)短暂增加,且温度升高3°C对响应持续时间没有影响。[CO₂]对生长的刺激是通过净同化率(nar)的持续增加来实现的,尽管由于单位叶质量的叶面积(sla)下降,叶面积比(lar)同时降低抵消了对rgr的潜在益处。对温度(T°C)的响应通常与对[CO₂]的响应相反。例如,T°C通过增加sla来提高lar,这而非对nar的任何影响是刺激rgr的主要因素。对[CO₂]效应的异速生长分析表明,在各个收获期观察到的分配变化完全是由于与植物大小相关的变化所致。相比之下,升高的温度(T°C)对叶片分配模式有直接影响,在生长初期叶面积相对于整株植物质量的扩展增加,随后生物量从叶片向植物其他部位的分配增加。在升高的[CO₂]或T°C条件下,根与地上部之间的异速生长关系均未观察到变化。因此,我们得出结论,生物量分配和诸如sla等形态特征对[CO₂]相对不敏感,至少在全株水平分析时如此,并且在观察到变化时,这些变化是比较相同年龄但不同大小的植物的结果。