Climate
driven increases in global terrestrial net primary
production from 1982 to 1999
Ramakrishna R. Nemani1,5,*
Charles D. Keeling2
Hirofumi Hashimoto1
William M. Jolly1
Stephen C. Piper2
Compton J. Tucker3
Ranga B. Myneni4
Steven W. Running1
1School
of Forestry,
2Scripps
Institution of Oceanography,
3NASA/Goddard
Space Flight Center,
4Department
of Geography,
5 After
September 2003, NASA/Ames Research Center,
* To whom
correspondence should be addressed. E-mail: nemani@ntsg.umt.edu
Recent climatic changes have enhanced plant
growth in
northern mid and high latitudes. However, a comprehensive analysis of the
impact of global climatic changes on vegetation productivity has not before
been expressed in the context of variable limiting factors to plant
growth. We present a global
investigation of vegetation responses to climatic changes by analyzing 18 years
(1982-1999) of both climatic data and satellite observations of vegetation
activity. Our results indicate that global changes in climate have eased
several critical climatic constraints to plant growth, such that net primary
production increased 6% (3.4 Pg C/18 yr) globally. The largest increase was in
tropical ecosystems. Amazon rainforests accounted for 42% of the global NPP
increase, owing mainly to decreased cloud cover and the resulting increase in
solar radiation.
Between 1980 and 2000 the Earth experienced dramatic environmental
changes (1). It had two of
the warmest decades in the instrumental record (1980s and 1990s), had three
intense and persistent El Niņo events (1982-1983, 1987-1988 and 1997-1998), and
saw significant changes in tropical cloudiness (2) and monsoon dynamics (3). Meanwhile,
atmospheric CO2 levels increased by 9% (337 to 369 ppm) and human
population increased by 37% (4.45x109 to 6.08 x109). Changes in terrestrial net primary production
(NPP) integrate these and other climatic, ecological,
geochemical, and human influences on the biosphere. Several regional studies have reported
increases in NPP (4-10), but a globally comprehensive analysis of the
impacts of climatic changes on NPP is lacking. For
the northern mid and high latitudes, these studies suggest that multiple
mechanisms (e.g., nitrogen deposition, CO2 fertilization, forest
regrowth and climatic changes) have promoted increases in NPP, while increases
in the tropics have been primarily attributed to CO2 fertilization.
Here we analyzed nearly two decades of recent global climatic data and
satellite observations of vegetative activity and
show that climatic changes have eased multiple climatic constraints to plant
growth, increasing NPP over large regions of the Earth.
Temperature, radiation,
and water interact to impose complex and varying limitations on vegetation
activity in different parts of the world (11). In order to provide a comprehensive interpretation
of climate change impacts on plant growth, we first constructed a map of the
relative contributions of climatic controls on global vegetation. We used long-term monthly climate statistics
to build simple bioclimatic indices (12). From these indices, we
estimated that water availability most strongly limits vegetation growth over
40% of the Earth's vegetated surface, temperature limits 33%, and radiation 27%
(Fig.1A). These factors tend to be co-limiting. For example, cold winter temperatures and
cloudy summers limit high latitude Eurasian vegetation, while cold winters and
dry summers limit vegetation in western
We have estimated the trends in these growth limiting climate factors
from 1982 to 1999 using daily reanalysis data from the
Most of the observed climatic changes have been in the direction of
reducing climatic constraints to plant growth. To quantify this effect, we used
a biome specific production efficiency model (PEM) (12, 17) that combines monthly estimates of satellite-derived
vegetation properties with daily NCEP climate data to estimate monthly and
annual NPP at 0.5 o x 0.5o resolution. The satellite-derived vegetation properties
used were: the fraction of absorbed
photosynthetically active radiation (FPAR) and leaf area index (LAI) derived
from remotely sensed Normalized Difference Vegetation Index (NDVI) and a biome
map (18, 19). The PEM that we used is similar in logic to other
PEMs (8, 13, 20) but has parameters derived from field studies and a
global ecosystem process model (17,
21). To
account for differences in satellite data processing, we used two independent
datasets of LAI and FPAR derived from Global Inventory Monitoring and Modelling
Studies (GIMMS) (22) and Pathfinder AVHRR Land (PAL) (23) NDVI data sets (12)
From 1982 to 1999, modelled NPP increased the most (6.5%) in water and
radiation limited regions followed by temperature and radiation limited regions
(5.7%), and temperature and water limited regions (5.4%). NPP increased significantly (p<0.01) over
25% of the global vegetated area, with a mean rate of 6.3 gC/m2/y,
and decreased significantly over only 7%, with a mean rate of 4.2 gC/m2/y
(Fig. 2).
To isolate the role of climate from other mechanisms that could enhance
carbon sequestration such as CO2 fertilization, nitrogen deposition
and forest re-growth, and to remove spurious trends in satellite data due to
residual sensor calibration effects, we alternately estimated NPP by assuming constant vegetation (1982-1999 monthly
average FPAR and LAI) but changing climate and by assuming constant climate (average daily climate from 1982-1999) with
changing vegetation. Changes in climate (with constant vegetation) directly
contributed nearly 40% of the total increase in NPP from 1982 to 1996 (Table
S2). Changes in vegetation (with
constant climate) over the same period contributed about 60% of total NPP
increase, possibly as a result of climate-vegetation feedbacks, changes in land
use and growth stimulation from other mechanisms. Over parts of
Globally, NPP increased (Fig. 3) by 6.17%, 3.42 PgC/18yr (p<0.001), between 1982 and 1999. Ecosystems in all tropical regions and those
in the high latitudes of the Northern Hemisphere accounted for 80% of the
increase. Although terrestrial NPP only
accounts for part of the biosphere-atmosphere carbon exchange, interannual
variations of NPP are negatively correlated with global increases in
atmospheric CO2 growth rate (r = 0.70, p<0.001). NPP anomalies attributable to climate alone
(calculated with constant vegetation) explained nearly the same amount of
variation in the NPP-CO2 relationship (r = 0.71, p<0.001),
indicating that climatic variability over land exerts a strong control over the
variation in atmospheric CO2 (Fig. S9).
NPP responded differentially with respect to latitude to major climatic
events such as El Niņo and volcanic eruptions (Fig. 4). Globally, NPP declined
during all three major El Niņo events with corresponding increases in global CO2
growth rate (24). El Niņo
events dominate tropical NPP variability, which has the highest association with global
CO2 growth rate (r = 0.75, p<0.001). This response is likely due to NPP and soil respiration
being more tightly coupled in tropical climates compared to ecosystems in other
latitudes. Soil carbon residence times range from less than four years in hot,
wet tropical areas to greater than 1000 years in cold boreal or dry desert
conditions (25). A strong decline in NPP following the
An
increase in NPP of only 0.2% per ppm increase in CO2, could explain
all of the estimated global NPP increase of 6.17%/18yr and is within the range
of experimental evidence (27). However, NPP increased by more than 1%/yr in
Quantifying
trends in NPP, as we have sought to do here, is necessary, but not sufficient
for understanding land surface net ecosystem exchange of CO2. The
same climatic changes that have caused decadal-scale increases in NPP can also
change above and below ground carbon allocation, decomposition rates,
disturbance regimes and other processes that cycle carbon between terrestrial
ecosystems and the atmosphere. Atmospheric CO2 inversion models show
that northern mid-latitude ecosystems have recently been consistently large
carbon sinks, and the tropics are either neutral or small sources, albeit with
high uncertainty (1). Our satellite-based estimates of NPP, on the other
hand, show significant growth stimulation in both the tropics and the northern
high latitude ecosystems. Assuming that carbon emissions, including those from
biomass burning and land use changes, are properly accounted for in the
atmospheric inversions, this spatial discrepancy means that respiration as well
as NPP is a major driver of terrestrial carbon sink dynamics. Global climate
models project, in response to increasing greenhouse gases, an intensified
hydrologic cycle altering the patterns of temperature, humidity, cloud cover
and rainfall (1). Our ability
to predict the future of terrestrial ecosystems is contingent upon how well we
can interpret such changes in the context of multiple limiting factors to
biogeochemical cycling.
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reviewers and Alisa Keyser for helpful comments. This work was supported
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support from NSF (ATM-01-20527), DOE (DE-FG03-95ER62075) and NASA (NAG5-11217).

Figure 1: Geographic distribution of
potential climatic constraints to plant growth derived from long-term climate
statistics (A), and recent climatic changes, estimated from reanalysis data
from 1982 to 1999, over the growing season of average temperature (B), of vapor
pressure deficit, reductions in VPD are indicative of increased water
availability (C), and of solar radiation (D). Growing season is defined as
those months with 1982-1999 average air temperatures above zero oC.

Fig. 2: Spatial
distribution of linear trends in estimated NPP from 1982 to 1999. NPP was
calculated using mean FPAR and LAI derived from GIMMS and PAL data sets.
Fig. 3:
Interannual variations from 1982 to 1999 in global NPP in relation to atmospheric
CO2 growth rate. Trends in global NPP anomalies are shown for GIMMS
(solid blue), PAL (dashed blue) and their average in green. CO2
growth rate (inverted) is shown in red.
Growth rates in ppm were converted to Pg of carbon using a conversion
factor of 2.12 Pg per ppm (24)). Mean NPP was
54.5 PgC/yr. Mean CO2 growth
rate was 3.2 Pg C/yr. A multivariate
ENSO index (MEI (31)) is shown in grey scale, where darker shades
represent higher MEI values.

Fig. 4: Interannual
variations in NPP distributed by latitudinal zones. Zonal NPP anomalies in each
zone are shown for GIMMS (solid blue), PAL (dashed blue), and their average in
green.