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Elevated CO2 and conifer roots: effects on growth, life span and turnover
- DAVID T. TINGEY, DONALD L. PHILLIPS, MARK G. JOHNSON
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- Journal:
- The New Phytologist / Volume 147 / Issue 1 / July 2000
- Published online by Cambridge University Press:
- 01 July 2000, pp. 87-103
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- July 2000
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Elevated CO2 increases root growth and fine (diam. [les ]2 mm) root growth across a range of species and experimental conditions. However, there is no clear evidence that elevated CO2 changes the proportion of C allocated to root biomass, measured as either the root[ratio ]shoot ratio or the fine root[ratio ]needle ratio. Elevated CO2 tends to increase mycorrhizal infection, colonization and the amount of extramatrical hyphae, supporting their key role in aiding the plant to more intensively exploit soil resources, providing a route for increased C sequestration. Only two studies have determined the effects of elevated CO2 on conifer fine-root life span, and there is no clear trend. Elevated CO2 increases the absolute fine-root turnover rates; however, the standing crop root biomass is also greater, and the effect of elevated CO2 on relative turnover rates (turnover[ratio ]biomass) ranges from an increase to a decrease. At the ecosystem level these changes could lead to increased C storage in roots. Increased fine-root production coupled with increased absolute turnover rates could also lead to increases in soil organic C as greater amounts of fine roots die and decompose. Although CO2 can stimulate fine-root growth, it is not known if this stimulation persists over time. Modeling studies suggest that a doubling of the atmospheric CO2 concentration initially increases biomass, but this stimulation declines with the response to elevated CO2 because increases in assimilation are not matched by increases in nutrient supply.
The potential effects of nitrogen deposition on fine-root production in forest ecosystems
- KNUTE J. NADELHOFFER
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- Journal:
- The New Phytologist / Volume 147 / Issue 1 / July 2000
- Published online by Cambridge University Press:
- 01 July 2000, pp. 131-139
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- July 2000
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Temperate forests are recipients of anthropogenic nitrogen (N) deposition. Because growth in these ecosystems is often limited by N availability, elevated N inputs from the atmosphere can influence above- and belowground production in forests. Although fine-root production is the largest component of belowground production in forests, it is unclear whether or how increases in Navailability to forest trees accompanying increased N deposition might influence fine-root growth. Uncertainties as to how fine-root dynamics (i.e. production and turnover) vary in relation to soil N availability contribute to this problem. Although fine-root biomass typically decreases along soil N availability gradients in forests, it is unclear whether fine-root production and turnover also decrease along these gradients. Here, four possible relationships between fine-root turnover, fine-root production, and forest soil N availability are evaluated to develop a general hypothesis about changes in rooting dynamics that might accompany increases in N deposition. The four possible relationships are as follows. (1) Fine-root turnover rates do not systematically change with N availability in forest soils. If this is true, then fine-root production rates decrease with fine-root biomass in relation to soil N availability, and increased N deposition could lead to decreased fine-root production in forests. (2) Decreases in photosynthate allocation belowground along N availability gradients will function to slow fine-root turnover (or increase life span) as N availability increases with N deposition, thereby dramatically decreasing fine-root production. (3) Fine-root production might increase with N availability even though fine-root biomass typically decreases with N availability. This could occur if fine-root metabolism and turnover increase (life span decreases) with soil N supply. Increases in fine-root production accompanying increases in N availability, if large enough, could result in constant proportions of forest production being allocated to fine roots as soil N availability increases with N deposition. (4) Although fine-root turnover and production might both increase as N becomes more available to tree roots, the proportional allocation of total primary production to fine roots could decrease. Identifying the most likely of these four possibilities requires intersite comparisons of forest root dynamics along gradients of soil N availability and N deposition. Collective results of studies that use sequential sampling of fine-root biomass to estimate production suggest that fine-root turnover and production either; do not vary systematically, or that they decrease as N availability increases. By contrast, studies using ecosystem C or N budgets suggest that fine-root turnover and production both increase with N availability and that similar increases might be expected with elevated N deposition. It is argued here that assumptions underlying most biomass-based estimates of fine-root production are more suspect than are assumptions underlying element budget-based estimates. If so, it is likely that N deposition will function to decrease forest fine-root biomass but to stimulate fine-root turnover and production. However, increases in fine- root turnover and production could eventually decrease if chronically elevated N deposition leads to forest stand mortality.
Global patterns of root turnover for terrestrial ecosystems
- RICHARD A. GILL, ROBERT B. JACKSON
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- Journal:
- The New Phytologist / Volume 147 / Issue 1 / July 2000
- Published online by Cambridge University Press:
- 01 July 2000, pp. 13-31
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- July 2000
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Root turnover is a critical component of ecosystem nutrient dynamics and carbon sequestration and is also an important sink for plant primary productivity. We tested global controls on root turnover across climatic gradients and for plant functional groups by using a database of 190 published studies. Root turnover rates increased exponentially with mean annual temperature for fine roots of grasslands (r2 = 0.48) and forests (r2 = 0.17) and for total root biomass in shrublands (r2 = 0.55). On the basis of the best-fit exponential model, the Q10 for root turnover was 1.4 for forest small diameter roots (5 mm or less), 1.6 for grassland fine roots, and 1.9 for shrublands. Surprisingly, after accounting for temperature, there was no such global relationship between precipitation and root turnover. The slowest average turnover rates were observed for entire tree root systems (10% annually), followed by 34% for shrubland total roots, 53% for grassland fine roots, 55% for wetland fine roots, and 56% for forest fine roots. Root turnover decreased from tropical to high-latitude systems for all plant functional groups. To test whether global relationships can be used to predict interannual variability in root turnover, we evaluated 14 yr of published root turnover data from a shortgrass steppe site in northeastern Colorado, USA. At this site there was no correlation between interannual variability in mean annual temperature and root turnover. Rather, turnover was positively correlated with the ratio of growing season precipitation and maximum monthly temperature (r2 = 0.61). We conclude that there are global patterns in rates of root turnover between plant groups and across climatic gradients but that these patterns cannot always be used for the successful prediction of the relationship of root turnover to climate change at a particular site.
Building roots in a changing environment: implications for root longevity
- D. M. EISSENSTAT, C. E. WELLS, R. D. YANAI, J. L. WHITBECK
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- Journal:
- The New Phytologist / Volume 147 / Issue 1 / July 2000
- Published online by Cambridge University Press:
- 01 July 2000, pp. 33-42
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- July 2000
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Root turnover is important to the global carbon budget as well as to nutrient cycling in ecosystems and to the success of individual plants. Our ability to predict the effects of environmental change on root turnover is limited by the difficulty of measuring root dynamics, but emerging evidence suggests that roots, like leaves, possess suites of interrelated traits that are linked to their life span. In graminoids, high tissue density has been linked to increased root longevity. Other studies have found root longevity to be positively correlated with mycorrhizal colonization and negatively correlated with nitrogen concentration, root maintenance respiration and specific root length. Among fruit trees, apple roots (which are of relatively small diameter, low tissue density and have little lignification of the exodermis) have much shorter life spans than the roots of citrus, which have opposite traits. Likewise, within the branched network of the fine root system, the finest roots with no daughter roots tend to have higher N concentrations, faster maintenance respiration, higher specific root length and shorter life spans than secondary and tertiary roots that bear daughter roots. Mycorrhizal colonization can enhance root longevity by diverse mechanisms, including enhanced tolerance of drying soil and enhanced defence against root pathogens. Many variables involved in building roots might affect root longevity, including root diameter, tissue density, N concentration, mycorrhizal fungal colonization and accumulation of secondary phenolic compounds. These root traits are highly plastic and are strongly affected by resource supply (CO2, N, P and water). Therefore the response of root longevity to altered resource availability associated with climate change can be estimated by considering how changes in resource availability affect root construction and physiology. A cost–benefit approach to predicting root longevity assumes that a plant maintains a root only until the efficiency of resource acquisition is maximized. Using an efficiency model, we show that reduced tissue Nconcentration and reduced root maintenance respiration, both of which are predicted to result from elevated CO2, should lead to slightly longer root life spans. Complex interactions with soil biota and shifts in plant defences against root herbivory and parasitism, which are not included in the present efficiency model, might alter the effects of future climate change on root longevity in unpredicted ways.
Dynamics of root systems in native grasslands: effects of elevated atmospheric CO2
- J. A. ARNONE, J. G. ZALLER, E. M. SPEHN, P. A. NIKLAUS, C. E. WELLS, C. KÖRNER
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- Journal:
- The New Phytologist / Volume 147 / Issue 1 / July 2000
- Published online by Cambridge University Press:
- 01 July 2000, pp. 73-85
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- July 2000
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The objectives of this paper were to review the literature on the responses of root systems to elevated CO2 in intact, native grassland ecosystems, and to present the results from a 2-yr study of root production and mortality in an intact calcareous grassland in Switzerland. Previous work in intact native grassland systems has revealed that significant stimulation of the size of root systems (biomass, length density or root number) is not a universal response to elevated CO2. Of the 12 studies reviewed, seven showed little or no change in root-system size under elevated CO2, while five showed marked increases (average increase 38%). Insufficient data are available on the effects of elevated CO2 on root production, mortality and life span to allow generalization about effects. The diversity of experimental techniques employed in these native grassland studies also makes generalization difficult. In the present study, root production and mortality were monitored in situ in a species-rich calcareous grassland community using minirhizotrons in order to test the hypothesis that an increase in these two measures would help explain the increase in net ecosystem CO2 uptake (net ecosystem exchange) previously observed under elevated CO2 at this site (600 vs 350 μl CO2 l−1; eight 1.2-m2 experimental plots per CO2 level using the screen-aided CO2 control method). However, results from the first 2 yr showed no difference in overall root production or mortality in the top 18 cm of soil, where 80–90% of the roots occur. Elevated CO2 was associated with an upward shift in root length density: under elevated CO2 a greater proportion of roots were found in the upper 0–6-cm soil layer, and a lower proportion of roots in the lower 12–18 cm, than under ambient CO2. Elevated CO2 was also associated with an increase in root survival probability (RSP; e.g. for roots still alive 280 d after they were produced under ambient CO2, RSP = 0.30; elevated CO2, RSP = 0.56) and an increase (48%) in median root life span in the deepest (12–18 cm) soil layer. The factors driving changes in root distribution and longevity with depth under elevated CO2 were not clear, but might have been related to increases in soil moisture under elevated CO2 interacting with vertical patterns in soil temperatures. Thus extra CO2 taken up in this grassland ecosystem during the growing season under elevated CO2 could not be explained by changes in root production and mortality. However, C and nutrient cycling might be shifted closer to the soil surface, which could potentially have a substantial effect on the activities of soil heterotrophic organisms as CO2 levels rise.
Spatial and temporal deployment of crop roots in CO2-enriched environments
- SETH G. PRITCHARD, HUGO H. ROGERS
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- Journal:
- The New Phytologist / Volume 147 / Issue 1 / July 2000
- Published online by Cambridge University Press:
- 01 July 2000, pp. 55-71
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- July 2000
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Growth of crops in CO2-enriched atmospheres typically results in significant changes in root growth and development. Increased root carbohydrates stimulate root growth either directly (functioning as substrates) or indirectly (functioning as signal molecules) by enhancing cell division or cell expansion, or both. Although highly variable, the literature suggests that, generally, initiation and stimulation of lateral roots is favored over the elongation of primary roots, leading to more highly branched, shallower root systems. Such architectural shifts can render root systems less efficient, perhaps contributing to the lower specific root activities often reported. Allocation of carbon (C) to roots fluctuates through the life of the plant; root functional and growth responses should therefore not be viewed as static. In annual crops, C allocation to belowground processes changes as vegetative growth switches to reproduction and maturation. Reductions in C allocation to roots over time might cause temporal shifts in root deployment, perhaps affecting root demography. However, significant changes in root turnover (defined here as root flux or mortality relative to total root pool size) as a result of decreased root longevities in crop plants are unlikely. Consideration of changing C allocation to roots, a more thorough understanding of the mechanistic controls on root longevity, and a better characterization of the rooting habits (life histories) of different crop species will further our understanding of how increasing atmospheric [CO2] will affect root demography. This knowledge will lead the way toward a more thorough understanding of the linkage of atmosphere with belowground plant function and also that of plant function with soil biology and structure. Ultimately, successful modeling of global C and nitrogen (N) cycles will require empirical data concerning spatial and temporal deployment of roots for a range of crop species grown under different agricultural management systems.
Root production and turnover and carbon budgets of two contrasting grasslands under ambient and elevated atmospheric carbon dioxide concentrations
- A. H. FITTER, J. D. GRAVES, J. WOLFENDEN, G. K. SELF, T. K. BROWN, D. BOGIE, T. A. MANSFIELD
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- Journal:
- The New Phytologist / Volume 137 / Issue 2 / October 1997
- Published online by Cambridge University Press:
- 01 October 1997, pp. 247-255
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- October 1997
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Monoliths of two contrasting vegetation types, a species-rich grassland on a brown earth soil over limestone and a species-poor community on a peaty gley, were transferred to solardomes and grown under ambient (350 μl l−1) and elevated (600 μl l−1) CO2 for 2 yr. Shoot biomass was unaltered but root biomass increased by 40–50% under elevated CO2. Root production was increased by elevated CO2 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 CO2 in the peat soil, but there was only a small effect on turnover in the limestone soil. Elevated CO2 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 CO2. 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. 0·2 kg C m−2 yr−1 (=2 t ha yr−1) under elevated CO2.