We have measured the rates of root production and death and of root respiration in situ under two grasslands along an altitudinal gradient in the northern Pennines, UK, represented by a lowland site at 171 m in an agricultural setting, and three upland sites between 480 and 845 m. One grassland was dominated by Festuca ovina and was on a brown earth soil; the other was dominated by Juncus squarrosus and Nardus stricta and occurred on a peaty gley. The natural altitudinal gradient was extended by transplantation. Although root biomass and root production (estimated using minirhizotrons) both showed pronounced seasonal peaks, there was no simple altitudinal gradient in either variable, and neither root production nor root death rate was a simple function of altitude. Increased root accumulation in summer was a function of change in the length of the growing season, not of soil temperature. Root populations in winter were similar at all sites, showing that increased production at some sites was accompanied by increased turnover, a conclusion confirmed by cohort analyses. Respiration rate, measured in the field by extracting roots and measuring respiration at field temperature in an incubator, was unrelated to temperature. The temperature sensitivity of respiration (expressed as the slope of a plot of log respiration rate against temperature) showed no simple seasonal or altitudinal pattern. Both root growth (under Festuca) and respiration rate were, however, closely related to radiation fluxes, averaged over the previous 10 days for growth and 2 days for respiration. The temperature sensitivity of respiration was a function of soil temperature at the time of measurement. These results show that root growth and the consequent input of carbon to soil in these communities is controlled by radiation flux not temperature, and that plants growing in these upland environments may acclimate strongly to low temperatures. Most carbon cycle models assume that carbon fluxes to soil are powerfully influenced by temperature, but that assumption is based largely on short-term studies and must be reassessed.

Sl'MMARYMonoliths of two contrasting vegetation types, a species-rich grassland on a brown earth soil over limestone and a species-poor community on a peat>' gley, were transferred to solardomes and grown under ambient (350 /d 1"') and elevated (600//I i"^) CO^ for 2 yr. Shoot biomass was unaltered but root biomass increased by 40-50% under elevated CO^. Root production was increased by elevated CO^ in the peat soil, measured both as instantaneous and cumulative rates, but only the latter measure was increased in the limestone soil. Root growth was stimulated more at 6 cm depth than at 10 cm in the limestone soil. Turnover was faster under elevated COî n the peat soil, but there was only a small effect on turnover in the limestone soil. Elevated CO^ reduced nitrogen concentration in roots and might have increased mycorrhizal colonization. Respiration rate was correlated with N concentration, and was therefore lower in roots grown at elevated CO^-Estimates of the C budget of the two communities, based upon root production and on net C uptake, suggest that C sequestration in the peat soil increases by c. 02 kg C m"^ yr"^ (= 2 t ha yr"^) under elevated CO^.

An essential component of an understanding of carbon flux is the quantification of movement through the root carbon pool. Although estimates have been made using radiocarbon, the use of minirhizotrons provides a direct measurement of rates of root birth and death. We have measured root demographic parameters under a semi-natural grassland and for wheat. The grassland was studied along a natural altitudinal gradient in northern England, and similar turf from the site was grown in elevated CO2 in solardomes. Root biomass was enhanced under elevated CO2. Root birth and death rates were both increased to a similar extent in elevated CO2, so that the throughput of carbon was greater than in ambient CO2, but root half-lives were shorter under elevated CO2 only under a JuncuslNardus sward on a peaty gley soil, and not under a Festuca turf on a brown earth soil. In a separate experiment, wheat also responded to elevated CO2 by increased root production, and there was a marked shift towards surface rooting: root development at a depth of 80-85 cm was both reduced and delayed. In conjunction with published results for trees, these data suggest that the impact of elevated CO2 will be system-dependent, affecting the spatio-temporal pattern of root growth in some ecosystems and the rate of turnover in others. Turrnover is also sensitive to temperature, soil fertility and other environmental variables, all of which are likely to change in tandem with atmospheric CO2 concentrations. Differences in turnover and time and location of rhizodeposition may have a large effect on rates of carbon cycling.

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