Woody-Tissue Respiration for Simarouba Amara and Minquartia Guianensis, Two Tropical Wet Forest Trees with Different Growth Habits
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Oecologia (1994) 100:213-220 9Springer-Verlag 1994 Michael G. Ryan 9Robert M. Hubbard Deborah A. Clark 9Robert L. Sanford, Jr. Woody-tissue respiration for Simarouba amara and Minquartia guianensis, two tropical wet forest trees with different growth habits Received: 9 February 1994 / Accepted: 26 July 1994 Abstract We measured CO 2 etflux from stems of two maintenance respiration was 54% of the total CO2 efflux tropical wet forest trees, both found in the canopy, but for Simarouba and 82% for Minquartia. For our sam- with very different growth habits. The species were ple, sapwood volume averaged 23% of stem volume Simarouba amara, a fast-growing species associated when weighted by tree size, or 40% with no size weight- with gaps in old-growth forest and abundant in sec- ing. Using these fractions, and a published estimate of ondary forest, and Minquartia guianensis, a slow-grow- aboveground dry-matter production, we estimate the ing species tolerant of low-light conditions in annual cost of woody tissue respiration for primary old-growth forest. Per unit of bole surface, CO2 efflux forest at La Selva to be 220 or 350 gCm-2year -1, averaged 1.24 gmol m -2 s-1 for Simarouba and depending on the assumed sapwood volume. These 0.83gmolm-2s -1 for Minquartia. C02 elttux was costs are estimated to be less than 13% of the gross highly correlated with annual wood production (r 2= production for the forest. 0.65), but only weakly correlated with stem diameter (r 2 = 0.22). We also partitioned the CO2 efttux into the Key words Woody tissue respiration" Maintenance functional components of construction and mainte- respiration 9Tropical wet forest trees 9Carbon balance nance respiration. Construction respiration was esti- mated from annual stem dry matter production and maintenance respiration by subtracting construction respiration from the instantaneous CO2 flux. Estimated Introduction maintenance respiration was linearly related to sap- wood volume (39.6 gmol m -3 s -1 at 24.6 ~ C, r 2 = 0.58), In forest ecosystems, autotrophic respiration may con- with no difference in the rate for the two species. strain productivity (Ryan 1991a) because forests con- Maintenance respiration per unit of sapwood volume tain much biomass in foliage, woody tissue, and roots. for these tropical wet forest trees was roughly twice that For example, the percentage of gross primary produc- of temperate conifers. A model combining construc- tion used for plant respiration may differ for forests tion and maintenance respiration estimated CO2 very growing in different climates or in response to climate well for these species (r 2= 0.85). For our sample, change because respiration rates are sensitive to tem- perature (Waring and Schlesinger 1985). Also, stabi- lization and subsequent decline of net primary production as stands age may be caused by the M. G. Ryan ([]) ' R. M. Hubbard BDA Forest Service, increased costs of woody-tissue respiration (Yoda et al. Rocky Mountain Experiment Station, 1965; Whittaker and Woodwell 1967), although this 240 West Prospect Street, hypothesis has been recently disputed (Ryan and Fort Collins, CO 80526-2098, USA Waring 1992). Rigorous tests of how autotrophic res- D. A. Clark piration affects forest productivity are rare (Jarvis and Organization for Tropical Studies, Leverenz 1983; Sprugel and Benecke 1991; Ryan and Apartado 676,2050, Waring 1992) and limited to temperate forests. San Pedro, Costa Rica In forest ecosystems in the wet tropics, autotrophic R. L. Sanford Jr. respiration can consume more than 50% of gross pri- Department of Biology, University of Denver, mary production (Edwards et al. 1980). Because of the Denver, CO 80208, USA large amount of woody biomass in these ecosystems 214 and the warm temperatures under which they grow, a volume. We also calculate an annual budget for woody- large fraction of the autotrophic respiration has been tissue respiration and compare it with other estimates. attributed to woody tissues (Mfiller and Nielson 1965; Yoda 1967; Whitmore 1984). These respiration esti- mates were developed using stem or branch diameter (Yoda 1967) or surface area (Odum 1970) to scale Methods chamber measurements to the stand. However, direct Study area measurements such as these can be highly variable due to differences in growth rate and proportion of living We conducted our study at the La Selva biological station in Costa tissue in the organ being measured (Ryan 1990; Sprugel Rica (10 ~ 26' N, 83 ~ 59' W). This lowland forest reserve is located 1990). Therefore, diameter or surface area may be poor at the base of the continental divide on the coastal plain of the Sarapiqui River at an elevation of 50 m. Mean annual rainfall is scalars for deriving stand estimates (Ryan 1990). 3880 mm with a moderately dry season from late January through Recent studies (Ryan 1990; Sprugel 1990) have April (Sanford et al. 1994). Mean monthly temperature varies from shown that it is possible to separate respiration in 24.7~ C in January to 27.1~ C in August with a mean annual tem- woody tissue into the functional components of con- perature of 26.2 ~ C. A water balance diagram for the site indicates struction (respiration used to build dry matter) and a considerable excess of precipitation over potential evapotranspi- ration for every month except for February and March when val- maintenance (respiration used to maintain ion gradi- ues are about equal (Herrera 1985). Old growth vegetation at La ents, replace enzymes, and repair membranes). The Selva has been described as premontane tropical wet forest in the functional model recognizes that the energy from res- Holdridge life-zone system (Hartshorn 1983). A complete descrip- piration is used for different purposes, even though it tion of forest composition and soils is given in McDade et al. comes from the same biochemical pathways. Construc- (1994). tion respiration will vary with the amount of tissue CO2 flux measurements built and its chemical composition, while energy used for maintenance varies with temperature and perhaps We measured CO2 elflux from 18 stems of Simarouba amara Aubl. with nutrition, water, atmospheric CO2 concentration (Simaroubaceae), a fast-growing species that requires a high-light ([CO2]), and pollutants (Amthor 1989). Using the func- environment, and 18 stems of Minquartia guianensis Aubl. (Olacaceae), a slow-growing canopy species. Trees were selected tional model has furthered our understanding of pol- from a long-term study of stem growth and tree population dynam- lutant effects (Amthor and Cumming 1988), differences ics (Clark and Clark 1992), and were located in primary forest on in respiration costs between young and old forests upland sites. We selected trees with a wide variety of diameters (Ryan and Waring 1992), and the response of respira- (6-66 cm) and growth rates (0-2.1 cmyear -l) to help separate tion to atmospheric [CO2] (Reuveni and Gale 1985; maintenance and construction respiration. For our sample, stem diameter and diameter growth averaged 27.0 cm (SD 16.2) and Amthor 1991; Amthor et al. 1992; Wullschleger et al. 9.7 mm (SD 5.1) for Simarouba and 21.6 cm (SD 15.0) and 1.7 mm 1992). To our knowledge, maintenance respiration has (SD 1.7) for Minquartia. never been estimated for woody tissue of trees in the CO2 efflux was measured in October 1991 using an open wet tropics. system infra-red gas analyzer (LCA3, Analytical Develop- ment Company), with flow rates through the chamber of In practice, construction respiration has been relat- 273-342 gmol s -1. We used three different chambers varying in size ed to dry matter production (Ryan 1990; Sprugel 1990) from 41 to 62 cm2; chamber volume was less than 100 cm 3. and its chemical composition (Vertregt and Penning de Chambers were constructed of Lexan and a small fan stirred the Vries 1987; Williams et al. 1987), and maintenance res- air inside the chamber. Chambers were sealed to the bark surface piration to tissue nitrogen content (Ryan 1991a) and with neoprene gasket and attached to the stems with elastic cord over the exact point where diameters were taken for the growth (for woody tissue) to sapwood volume (Ryan 1990; study (1.5~4 m above the ground), Reference air was drawn through Sprugel 1990; Ryan et al. 1995). The use of the func- a 20-1 mixing chamber to maintain a stable reference COz concen- tional model of respiration can explain the variability tration. Flux data were recorded after the difference in [CO2] in chamber measurements and the derived respiration between reference and chamber had been stable for 3 min. We found no appreciable variation in CO2 efflux from different positions rates can be directly incorporated into process-based around the circumference of the bole or in measurements repeated models of forest growth [e.g., BIOMASS (McMurtrie within 1 h at the same location on the bole. Therefore, we sampled et al. 1990), HYBRID (Friend et al. 1993), Forest- most boles only once. CENTURY (Sanford et al. 1991)]. To successfully model the productivity of tropical Estimating construction and maintenance respiration forests and the response to climate change, we need In temperate forests, maintenance respiration for woody tissues has reliable models for respiration of the woody tissue of been estimated by measuring CO2 efftux after growth ceased in tropical forest trees. In this study, we measure stem res- autumn (Ryan 1990; Sprugel 1990; Sprugel and Benecke 1991). This piration for two very different trees found in the wet approach could not be used where stem growth continues through- tropics. We estimate maintenance respiration by sub- out the year. Therefore, we estimated maintenance respiration by subtracting construction respiration (estimated from stem growth tracting construction respiration (estimated using stem and wood density) from measured CO2 efftux.