Recent global-scale analyses indicate that climate variability affects net carbon storage but regard temperature and precipitation to be the main contributors. Seasonal and interannual variation in light availability may also limit CO 2 uptake. As an experimental test of light limitation by cloud cover during tropical rainy seasons and by the unusually heavy cloud cover associated with La Niñ a, we installed high-intensity lamps above the forest canopy to augment light for Luehea seemannii, a tropical canopy tree species, during cloudy periods of 1999 -2000. Light augmentation only partially compensated for the reduction in photosynthetic photon flux density caused by clouds. Nonetheless, leaves acclimated to the augmented irradiance, and photosynthesis, vegetative growth, and reproduction increased significantly. Light, rather than water, temperature, or leaf nitrogen, was the primary factor limiting CO 2 uptake during the rainy season.El Niñ o ͉ canopy ͉ photosynthesis ͉ light limitation W e ask whether year-to-year variation in cloud cover and light intensity limits photosynthesis and carbon uptake by a widespread tropical tree. This question is important for several reasons. Heavy tropical cloud cover can reduce irradiance and limit photosynthesis by fully exposed sun leaves (1-3). Recent eddy covariance studies indicate that day-to-day variation in cloud cover and irradiance also affects net carbon uptake by forests (4-6). Tropical cloud cover varies seasonally (7,8) and interannually with fluctuations in the El Niño Southern Oscillation (9). Tropical cloud cover also underwent strong longitudinal and latitudinal redistributions during the 1990s (10), and global solar irradiance reaching the Earth's surface has declined by 2.7% per decade since the 1950s, probably as a consequence of increases in anthropogenic particulates and cloud cover (11). Finally, tropical forests account for 32-36% of global terrestrial net primary production (12, 13). The implications for global carbon uptake are significant if carbon uptake by tropical trees is indeed limited by year-to-year variation in cloud cover and irradiance.To determine whether natural year-to-year variation in cloud cover limited carbon gain, we supplemented light levels artificially whenever cloud cover reduced photosynthetic photon flux density (PPFD; 400-700 nm) by 700 mol⅐m Ϫ2 ⅐s
Ϫ1for two replicate adult canopy trees in a tropical forest located in the Republic of Panama. Supplemental illumination compensated only partially for natural reductions in PPFD. This experimental manipulation was maintained for 2 years, including 1999, the particularly cloudy La Niña year. We studied the response to augmented light of leaf-level photosynthesis, branch-level sap flow, leaf-and branch-level carbohydrate storage, branch extension growth, and fruit production.
Materials and MethodsSite, Species, and Treatments. The study was conducted at the Parque Natural Metropolitano (PNM; lat. 9°04ЈN, long. 79°23ЈW), a 256-hectare natural reserve located near Panama City, Rep...