TerrestrialTerrestrial carboncarbon sinkssinks –– uncertainuncertain Lowland rainforest, Borneo (F Lanting) © Mindenexplanations Pictures/FLPA R A Houghton Woods Hole Research Center, USA The carbon cycle is fundamental to life on Earth and it determines the amount of carbon Carbon is the basis of life on Earth. It is incorporated into amount of carbon in soils is large and variable, it is difficult to plants through photosynthesis, absorbed by animals through measure an annual carbon sink, even in a small, well-defined their food, present in the atmosphere as carbon dioxide, area. To measure a change globally is nearly impossible. locked into rock as limestone, and pressed into fossil fuels Given that terrestrial ecosystems are accumulating carbon in such as coal and oil. In the Jurassic age, there was much some regions and releasing it in others, it is very difficult to more carbon in the atmosphere than there is now. The carbon determine, directly, the magnitude of the terrestrial carbon taken up by plants on land and in the sea gradually, over sink. (See Box 1 for a description of the methods used to millennia, exceeded the amount released during decay, and measure or infer the magnitude of terrestrial sinks.) this ‘excess’ carbon became locked away as fossil fuels Direct measurement of changes in the concentrations of beneath the surface of the planet. In effect, life on Earth was carbon dioxide and oxygen in the atmosphere provides one acting as a ‘carbon sink’. For the last two centuries, this trend estimate of the global terrestrial sink (Table 1). Nearly half of has been drastically reversed as forests have been reduced the carbon released each year from fossil fuel combustion and fossil fuels have been burnt, meaning that more carbon accumulates in the atmosphere. The oceans and some has been released into the atmosphere than has been terrestrial ecosystems take up the rest. According to a recent absorbed. There is considerable concern that this is leading global analysis based on changes in the concentrations of directly to global warming. ‘Carbon sinks’ are now the focus of carbon dioxide and oxygen (Plattner et al., in review), the considerable study. world’s terrestrial ecosystems (vegetation and soils) were a Although the oceans are currently the greatest carbon sink, small net sink during the 1980s and 1990s. terrestrial carbon sinks are also important. The greatest Independent of these estimates of a net terrestrial sink, terrestrial carbon sinks occur in young, growing forests, changes in land use (including deforestation – whether for because a hectare of trees holds up to 50 times more carbon settlements or agriculture – reforestation, and the manage- than a hectare of grasses or crops. Older forests and soils ment of forests and wood products) are calculated to have may also accumulate carbon but the rates are generally low released an average of 2.0 x 10 15 grams of carbon per year compared with the rate for young trees. Nevertheless, low (PgC yr-1), and slightly more during the 1990s (Houghton, in rates over large areas, including grasslands as well as press). These estimates include the releases of carbon from forests, can yield a sink that is globally large. Because the the oxidation of wood products and logging debris, as well as Biologist (2002) 49 (4) 155 Terrestrial carbon sinks Box 1. Methods that are used to measure or infer carbon sinks 1. Global budgets based on atmospheric data and models areas of a few km 2 , but is difficult to scale up to larger The CO2 concentration of air is measured weekly at a regions. Because net fluxes of carbon are generally small network of nearly 100 sites around the world. Using models relative to the gross daytime and night time fluxes, the of atmospheric transport, scientists calculate the surface approach is better suited for determining short-term sources and sinks of carbon that explain the observed varia- responses of photosynthesis and respiration to variations in tion in CO2 concentrations. This inverse modelling approach temperature and moisture than to determining annual can also be used to distinguish terrestrial vs. oceanic carbon carbon budgets. sinks when the atmospheric measurements include tempo- ral variations in O2 and in the isotopic composition of CO 2 5. Physiologically-based process models of ecosystems 13 (i.e., CO2; Table 5). Ecosystem- and global-scale models simulate changes in carbon storage as a result of environmentally induced 2. Global budgets based on models of oceanic carbon changes in photo- uptake synthesis, respiration, growth, litter fall and decomposition. Models of the oceans’ carbon cycle (including carbon chem- The advantage of such process-based models is that they can istry and oceanic circulation) can be used with temporal pat- be used to distinguish the effects of different factors on carbon terns of atmospheric CO2 and fossil fuel emissions to infer the storage. The disadvantage is that it is difficult to know whether net terrestrial sources and sinks of carbon. This approach they include all of the important processes. yields a global value (no regional distinctions), but can be applied over the last two centuries. CO2 concentrations 6. Carbon models based on changes in land use before 1957, when direct measurements began, are These models are based on the changes in vegetation and obtained from analysis of the air trapped in bubbles in soil that accompany a change in land use. When forests are Greenland and Antarctic glaciers. converted to agricultural land, for example, the amount of carbon in the vegetation and soil is reduced. When open 3. Regional carbon budgets constructed from forest invento- lands are afforested, carbon stocks increase. The rates of ries loss and increase vary with land use and type of ecosystem, Most of the developed nations have national forest invento- but they are documented for many systems. Most of the ries that monitor the volume of wood in forests. Changes in error in the calculated flux of carbon results from uncertain- volumes over time indicate sources or sinks of carbon. ties in rate of deforestation and wood harvest. Supplementary information can be used to infer associated changes in woody debris, wood products, and soil carbon – Estimates of carbon flux based on changes in land use can those components that are not directly measured in the inven- theoretically identify sources and sinks resulting from direct tories. human activity, but to do so requires baseline information on the rates of forest growth that would have existed in the 4. Stand-level direct measurement of CO 2 flux (from towers) absence of changes in CO2 , N, climate, or other Direct measurement of CO2 flux over an ecosystem is environmental variables. Such data are generally unavail- obtained by linking CO2 concentrations in air with the upward able. Nevertheless, the method, in combination with meth- or downward movement of that air (the eddy covariance ods that measure a net source or sink (methods 1–4 above), technique). The approach yields hourly estimates of flux for has the potential to distinguish sinks resulting from direct the accumulations of carbon in regrowing forests. Strangely, the atmosphere. These factors generally result indirectly from the difference between the net terrestrial sink and the emis - human activities. sions from land-use change suggests that there is a residual • Disturbance and recovery mechanisms, including both terrestrial sink, not well understood, that locked away as natural disturbances and the direct effects of changes in much as 3.0 PgC yr-1 during the last two decades (Table 1). land use and management. Disturbances affect the mortal- The exact magnitude, location and cause of this residual ity of forest stands and thus the age structure of forests; terrestrial sink are uncertain, but it is important to understand recovering forests are sinks for carbon. Different land the mechanisms responsible for at least two reasons. First, management strategies may also affect the amount of the Kyoto Protocol requires that sinks resulting directly from carbon stored in non-forest ecosystems. human activity (e.g., planting trees) be distinguished from sinks unrelated to direct human management (e.g., growth Physiological or metabolic mechanisms enhancement through CO2 fertilisation), so as to avoid ‘undue credits’. Second, some potential mechanisms are responsible for the current sink more likely to persist into the future than others are. Without The longest-standing physiological mechanism proposed for a greater understanding of the global carbon cycle, plans for the accumulation of carbon on land is CO2 fertilisation. remedial action will be, at best, reliant on some guesswork. Horticulturalists have long known that annual plants respond There is even the chance of inadvertently worsening the situ- to higher levels of CO2 with increased rates of growth. The ation by harming existing sinks. concentration of CO2 in greenhouses is often deliberately The mechanisms proposed as being responsible for terres- increased to make use of this effect. Similarly, experiments trial sinks fall into two broad categories (Table 2): have shown that most C3 plants (all trees, most crops, and • Physiological or metabolic factors that affect rates of photo- vegetation from cold regions) respond to elevated concentra- synthesis, respiration, growth, and decay. These factors tions of CO2 with increased rates of photosynthesis and include elevated concentrations of atmospheric growth. CO 2, increased availability of nutrients, and changes in Despite the observed stimulatory effects of CO2 on photo- temperature and rainfall, any of which could increase synthesis and plant growth, it is not clear that the effects will growth rates in forests – locking more carbon away from result in an accumulation of carbon (that is, a net sink) in 156 Biologist (2002) 49 (4) Terrestrial carbon sinks Table 1. The global carbon budget. Units are PgC yr- 2001).