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105
6
Missing Gaps in the Estimation
of the Carbon Gains Service from
Light Use Efficiency Models
A. J. Castro Martínez
University of Oklahoma, Oklahoma, USA; University of Almería, Spain
J. M. Paruelo
University of Buenos Aires, Argentine
D. Alcaraz-Segura
University of Granada, Spain; University of Almería, Spain
J. Cabello
University of Almería, Spain
M. Oyarzabal
University of Buenos Aires, Argentine
E. López-Carrique
University of Almería, Spain
CONTENTS
6.1 Introduction ................................................................................................ 106
6.2 Material and Methods ............................................................................... 107
6.3 Results ......................................................................................................... 108
6.3.1 Estimation Methods and LUE Units ........................................... 108
6.3.2 LUE Estimates across Organizational Levels and
Land Cover Types .......................................................................... 110
6.3.3 Time Interval of LUE Estimates ................................................... 112
6.4 Conclusions ................................................................................................. 115
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106 Earth Observation of Ecosystem Services
6.1 Introduction
The scientic community is being urged to invest more time and economic
resources to improve current estimates of global and regional carbon bud-
gets (Scurlock etal. 1999). Carbon gains are considered either as an interme-
diate service (Fisher etal. 2009) or as supports of provision and regulating
services (MA 2005). In addition, net primary production (NPP), an esti-
mate of ecosystem carbon gains, is often considered the most integrative
descriptor of ecosystem function (McNaughton etal. 1989). NPP estimates
are derived from biomass harvesting, ux tower measurements, remote
sensing, and model simulation (Ruimy etal. 1995; Sala etal. 2000; Stilletal.
2004). Biomass harvesting is expensive and not exempt from errors and
methodological problems. These methods are limited in their spatial and
temporal coverage. Given the linear relationship between the fraction of
solar radiation absorbed by vegetation and spectral vegetation indices
(Sellers etal. 1992), Monteith’s model (Monteith 1972) offers the possibil-
ity of estimating seasonal variation in carbon gains from remote sensing
data (Potter 1993). Monteith’s model states that carbon gains (Equation 6.1)
of vegetation cover are a function of the quantity of incoming photosyn-
thetically active radiation (PAR), the fraction of this radiation intercepted
by vegetation (fPAR), and the light use efciency (LUE; Still et al. 2004).
The ux estimated using the Monteith’s model included net and gross pri-
mary production and net ecosystem exchange (NEE) (Ruimy etal. 1999; see
Equations6.2 and 6.3).
NPP = PAR*fPAR*LUE (6.1)
GPP = PAR*f PAR*LU E (6 .2)
NEE = PAR*fPAR*LUE (6.3)
PAR can be directly measured using radiometers; fPAR can be esti-
matedfrom spectral indices such as the Normalized Difference Vegetation
Index(NDVI; Asrar etal. 1984) or the Enhanced Vegetation Index (EVI). The
relationship of fPAR-spectral indices may vary between land cover types,
but several authors have proposed different empirical relationships: (a) lin-
ear (Choudhury 1987); (b) nonlinear (Potter 1993; Sellers etal. 1994); and
Acknowledgments .............................................................................................. 115
Appendix 6.1 (Articles Reviewed from 1972 to 2007) ................................... 115
References .............................................................................................................121
K14591_C006.indd 106 28/08/13 9:11 PM
107Missing Gaps in the Estimation of the Carbon Gains Service
(c)a combination of both (Los etal. 2000). The LUE termhas a maximum
value comparable to the photosynthetic efciency or quantum yield at leaf
level under optimum conditions (Gower etal. 1999). However, low tempera-
tures and water and nutritional stress reduce LUE value (Field etal. 1995;
Gamon etal. 1995). Field etal. (1995) reported LUE differences from0.27gC/
MJ APAR for deserts to 0.70 g C/MJ APAR for tropical forests.
LUE was rst dened at the species level and mainly for crop species
(Andrade etal. 1993; Kiniry etal. 1998). The use of Monteith’s model as the
conceptual framework for remotely sensed estimates of aboveground net pri-
mary productivity (ANPP) (see Chapter5) requires a denition of LUE at the
ecosystem level (Ruimy etal. 1999; Sala etal. 2000; Fensholt etal. 2006). Often
a single xed value of approximately 1 g C/MJ APAR is used for a wide range
of spatiotemporal situations (Maselli etal. 2009). Several authors showed that
LUE varied in space (Field etal. 1995; Paruelo etal. 2004; Tongetal. 2008;
Garbulsky etal. 2010) and time (Nouvellon etal. 2000; Piñeiro et al. 2006)
and that the use of a single value may lead to substantial errors in regional
(Hilker etal. 2008) and global (Turner etal. 2002, 2003, 2005; Tong etal. 2008)
estimates of carbon gains.
Many factors affect the spatiotemporal patterns of LUE variation. Species
composition, plant structure and physiology, including leaf form and
RUBISCO (Ribulose-1,5-bisphosphate carboxylase oxygenase) content (Zhao
etal. 2007), and environmental factors (i.e., water stress, CO2 concentration,
temperature) modify LUE at the ecosystem level. Measuring LUE is not a
simple task. LUE can be estimated at different levels of organization (e.g.,
from individuals to ecosystems), using leaf-level estimates for single indi-
viduals or eddy covariance towers to derive LUE values at the ecosystem
level (Garbulsky etal. 2010).
LUE is the more uncertain parameter of the Monteith’s model since it is
not possible to measure it directly, and it depends on estimates of GPP/NPP/
NEE and absorbed radiation (Gower etal. 1999; Ruimy etal. 1999). This chap-
ter reviews the reported estimates of LUE, and the effect of the time inter-
val in LUE estimation at different organizational levels, from individuals
to ecosystems. We sought to answer the following questions: (1) How was
LUE estimated? (2) How did LUE differ across land cover types and levels
of organization? (3) How variable are LUE estimates according to the time
interval of estimation?
6.2 Material and Methods
We reviewed 125 articles from 1972 to 2007 containing the terms “light use
efciency” and “radiation use efciency,” but only 101 provided quantita-
tive LUE data (Appendix 6.1). The review included 65 different journals
K14591_C006.indd 107 28/08/13 9:11 PM
108 Earth Observation of Ecosystem Services
primarily in the eld of ecology (72% of total studies) and remote sensing
(22% of total studies). From the published studies, we developed a data-
base that included the organizational level, the ux estimated, LUE esti-
mates, and the geographical coordinates of the study site (Table6.1). We
characterized LUE values at three organizational levels: (1) “Individual”
referred to local-scale studies that estimated LUE based on individuals
of a single species; (2)“Single-species-dominated ecosystems” referred to
when the study focused on plots with one dominant species (e.g., NEE
of agroecosystem in eddy covariance ux tower with a footprint of 100
m2); and (3)“Multispecies-dominated ecosystems.” Assuming that 50% of
the dry biomass corresponds to carbon, LUE values were transformed to
the most common unit system: grams of carbon xed per megajoules of
absorbed PAR (g C/MJ APAR). To analyze the variability in LUE data, we
assigned each data to one category of Archibold’s (1995) classication of
terrestrial land cover types. We calculated the mean, maximum, and mini-
mum LUE values, as well as the deviation for each organizational level and
land cover type. Kruskal–Wallis tests were applied to detect signicant
differences in LUE estimates (n = 185) across organizational levels and
land cover types.
6.3 Results
6.3.1 Estimation Methods and LUE Units
LUE values were estimated using two main approaches. In the rst approach
(82% of total studies), LUE values were calculated at a local scale using the
Monteith’s equation and based on previous eld estimates of a carbon ux
(i.e., NPP or GPP). Here, the 20% of carbon ux estimates were derived from
CO2 ux between atmosphere and vegetation observations using eddyco-
variance techniques (Ruimy etal. 1995; Zhao etal. 2007); fPAR data were
calculated as a lineal function of satellite-derived NDVI in 44% of the total
data. The remaining studies used fPAR data reported from other studies or
compiled from direct measurements of the canopy. In most of the studies,
PAR was calculated by radiometers.
In the second approach (18% of total studies), LUE was estimated based
on correlative models with other variables such as leaf area index (LAI) or
the photochemical reectance index (PRI; Gu etal. 2002; Filella etal. 2004;
Graceet al. 2007) (see Chapter3). Here, LUE was also derived as a ratio
between the harvested biomass at the plot scale and the incoming APAR
throughout an entire year or a growing season.
Most of the articles reviewed (77% of total studies) offered quantitative
estimates of LUE. From these studies, we obtained 185 LUE values that
K14591_C006.indd 108 28/08/13 9:11 PM
109Missing Gaps in the Estimation of the Carbon Gains Service
TABLE6.1
Summary Sample of Light Use Efciency Data Reviewed for 1972–2007
Land
Cover
Types
Number
of Studies Locations LUE Units
Carbon Flux Model
(expressed as
percentage of total
studies within biome)
PHM 1 Alaska g C/MJ APAR (100%) Other*
CF 49 Durham, Canada,
Wisconsin,
Sweden
g C/MJ APAR Mol
C/mol photons
Moles CO2/
molPAR
(8%) GPP = APAR*LUE
(16%) NEE = APAR*LUE
(32%) NPP = APAR*LUE
(44%) Other*
TW 12 Canada, Europe,
EEUU
g C/MJ APAR
Mol C/mol
photons
Moles CO2/
molPAR
(8%) ANPP = APAR*LUE
(29%) NPP = APAR*LUE
(29%) Other*
Cr 38 Ireland, EEUU,
China, Italy,
Australia, United
Kingdom, South
Africa, India
g C/MJ
Kg (CO2/ha·h)/
(J/m2·sg)
(30%) NPP = APAR*LUE
(70%) Other*
TFE 65 Europe, EEUU,
Japan, China,
New Zealand
g C/MJ APAR
Mol CO2/
molAPAR
mmol CO2/
mmolphotons
(8%) GPP = APAR*LUE
(32%) NEE = APAR*LUE
(27%) NPP = APAR*LUE
(33%) Other*
ME 12 Spain, Italy, India,
EEUU
g C/MJ APAR
Mol C/mol APAR
(20%) NEE = APAR*LUE
(70%) NPP = APAR*LUE
(10%) Other*
TG 16 Canada, EEUU,
Argentina
g C/MJ APAR
g DM/MJ
(13%) NEE = APAR*LUE
(40%) NPP = APAR*LUE
(47%) Other*
TpF 4 Panama,
Colombia, EEUU
g C/MJ APAR
Kg(CO2/ha·h)/
(J/m2·sg)
(67%) GPP = APAR*LUE
(33%) NEE = APAR*LUE
TpS 9 Senegal, Argentina g C/MJ APAR
Mol C/mol APAR
(100%) Other*
AR 8 Sahara, Southern
Australia, Mali,
Mexico
g C/MJ APAR
gr DM/MJ
(17%) GPP = APAR*LUE
(50%) NPP = APAR*LUE
(33%) Other*
Note: Other* expresses (a) when the study did not specify the carbon ux model for LUE esti-
mation, (b) modications of Monteith’s model such as NASA-CASA model (i.e.,Carnegie-
Ames-Stanford Approach) simulates net primary productivity and the soil heterotrophic
respiration at regional to global scales or the TURC model for the estimation of the conti-
nental gross primary productivity and net primary productivity, or derived models by
Montetih’s approach based on the inclusion of other physiological parameters, and (c) a
constant LUE value. Archibold’s land cover type classication: PHM = polar and high
mountain tundra, CF = coniferous forests, TW = terrestrial wetlands, Cr = crops,
TFE= temperate forest ecosystems, ME = Mediterranean ecosystems, TG = temperate
grasslands, TpF = tropical forests, TpS = tropical savannas, and AR = arid regions.
K14591_C006.indd 109 28/08/13 9:11 PM
110 Earth Observation of Ecosystem Services
were originally expressed in four different units, including g C/MJ APAR
(65%), mol CO2/mol APAR (14%), g of dry matter/MJ APAR (9%), and mol
C/mol absorbed photons per minute (3%). After converting to g C/MJ
APAR, the average LUE value was 0.99 g C/MJ APAR (SD = 1.09), with
an absolute maximum of 8.2 g C/MJ APAR and an absolute minimum of
0.05gC/MJ APAR.
6.3.2 LUE Estimates across Organizational Levels and Land Cover Types
The number of studies varied among organizational levels and land cover
types (Figure 6.1). The multispecies-dominated ecosystems level was the
most commonly studied (62%) followed by single-species-dominated eco-
systems (19%) and individuals (19%). Temperate forests, coniferous forests,
and croplands were the most highly represented land cover types in the lit-
erature reviewed, and the least represented were polar and high mountain
tundra, tropical forests, and arid regions (Figure6.1). Croplands and temper-
ate forest ecosystems were the unique land cover types studied at all levels
of organization.
We found that the average LUE values at the individual level
(1.7gC/MJAPAR; SD = 1.6) were signicantly higher than at the multispe-
cies-dominated (0.8gC/MJ APAR; SD = 0.9) and single-species-dominated
Frequency (%)
40
30
20
10
0
PHM CF TW Cr TFE ME TG TpF TpS AR
1% 5%
0.5%
0.5%
2%
3%
17%
11%
9%
3%
5%
3%
2%
8%
5%
31%
17%
22%
6%
2%
20%
1%
Land cover types
Individuals (19%)
Single-species-dominated
ecosystems (19%)
Multispecies-dominated
ecosystems (62%)
FIGURE 6.1
Frequency of articles for land cover types at individual, single-species-dominated ecosystems,
and multispecies-dominated ecosystems levels. Total LUE values = 185. Archibold’s land cover
type classication: PHM = polar and high mountain tundra, CF = coniferous forests, TW =
terrestrial wetlands, Cr=crops, TFE = temperate forest ecosystems, ME = Mediterranean eco-
systems, TG= temperate grasslands, TpF = tropical forests, TpS = tropical savannas, and AR =
arid regions.
K14591_C006.indd 110 28/08/13 9:11 PM
111Missing Gaps in the Estimation of the Carbon Gains Service
(0.7g C/MJ APAR; SD = 0.4) ecosystems levels (Figure 6.2). Average LUE
values by land cover type exhibited signicant differences (Figure6.3). The
average LUE values varied between 2.20 g C/MJ APAR (SD = 1.67) in crop-
lands and 0.55gC/MJ APAR (SD = 0.23) in terrestrial wetlands. The maxi-
mum LUE was found in crops (8.20 g C/MJ APAR) and temperate grasslands
(5.20gC/MJ APAR), whereas the minimum LUE value was observed in trop-
ical savannas (0.05gC/MJ APAR) and temperate grasslands (0.06 g C/MJ
APAR)(Figure6.3).
At the individual level, signicant differences in LUE were observed
between tropical savannas, tropical forests, and crops (Figure 6.4). At the
single-species-dominated ecosystems level, coniferous forests and crops
were signicantly different in LUE values (Figure6.4). At the multispecies-
dominated ecosystems level, crops were signicantly different from conif-
erous forests, terrestrial wetlands, tropical forests, and Mediterranean
ecosystems. Temperate grasslands and tropical forest ecosystems were signif-
icantly different from coniferous forest, terrestrial wetlands, and temperate
forest ecosystems (Figure6.4).
Although LUE estimates were not available in the literature for all
organizational levels, we observed signicant differences within land
6
5
(116) (34) (35)
4
3
2
1
LUE (g C/MJ APAR)
0
Multispecies-
dominated ecosystems
Single-species-
dominated ecosystems Individuals
FIGURE 6.2
Box plot of light use efciency (LUE) values at multispecies-dominated ecosystems, single-
species- dominated ecosystems, and individual levels. The graphic explains t he minimum, rst
quartile (25%), median, mean, and third quartile (75%) of LUE values. The mean is displayed
with a +, and a black line corresponds to the median. The maximum LUE value at the indi-
vidual level was 8.2g C/MJ APAR; at the single-species-dominated ecosystems level, it was
2gC/MJ APAR; and at multispecies-dominated ecosystem level, it was 5.7 g C/MJ APAR. The
horizontal dotted line represents the total average of LUE values. The total LUE values per box
plot appear in brackets.
K14591_C006.indd 111 28/08/13 9:11 PM
112 Earth Observation of Ecosystem Services
cover types. Coniferous forests showed signicant differences between
single-species-dominated and multispecies-dominated ecosystems levels. In
temperate forest ecosystems, we observed signicant differences between
the individual and single- species-dominated ecosystems and multispecies-
dominated ecosystems levels (Figure6.4). In Mediterranean ecosystems, dif-
ferences were observed between the individual and multispecies-dominated
ecosystems levels. Crops, represented at all levels of organization, were the
only land cover type showing no signicant differences. Tropical forest eco-
systems with values for individual and multispecies-dominated ecosystems
levels did not exhibit signicant differences (Figure6.4).
6.3.3 Time Interval of LUE Estimates
LUE values signicantly differed according to the time interval of the
estimation (i.e., sampled within a day, season, or year). Annual and sea-
sonal estimates were obtained for all organizational levels. Daily LUE
estimates were only found in the literature at the multispecies-domi-
nated ecosystems level. Signicant differences between measurement
time intervals were detected at the single-species-dominated ecosystems
level between annual and seasonal LUE estimates. Kruskal–Wallis tests
did not reveal signicant differences in annual and seasonal estimates
at the individual level and for annual, seasonal, and daily estimates at
LUE (g C/MJ APAR)
(1)
0
PHM CF TW Cr TFE ME TG TpF TpS
AR
1
2
3
4
5
6
(41) (12) (32) (58) (10) (15) (2) (9) (5)
FIGURE 6.3
Box plot of light use efciency (LUE) values by land cover types. The graphic explains the
minimum, rst quartile (25%), median, mean, and third quartile (75%) of LUE values. The
mean is displayed with a +, and a black line corresponds to the median. Total LUE values =
185. Archibold’s land cover type classication: PHM = polar and high mountain tundra, CF =
coniferous forests, TW = terrestrial wetlands, Cr=crops, TFE = temperate forest ecosystems,
ME = Mediterranean ecosystems, TG = temperate grasslands, TpF = tropical forests, TpS =
tropical savannas, and AR = arid regions.
K14591_C006.indd 112 28/08/13 9:11 PM
113Missing Gaps in the Estimation of the Carbon Gains Service
the multispecies-dominated ecosystems level. However, we observed
signicant differences between annual and seasonal estimates at the
single- species-dominated ecosystems level. A comparison between the
time interval of the estimation and the organizational levels (Figure6.5)
revealed signicant differences between LUE obtained in annual and sea-
sonal observations. Annual and seasonal estimates were not signicantly
different at the multispecies-dominated and single-species-dominated
ecosystems levels, but we did detect signicant differences with the results
obtained at the individual level.
LUE (g C/MJ APAR) LUE (g C/MJ APAR) LUE (g C/MJ APAR)
6C
CA
cd
BCDH J
efg
g
gf
e
e
g
ef
ef
PHM
(1)
CF
(41)
TW
(12)
Cr
(32)
TFE
(58)
ME
(10)
TG
(15)
TpF
(2)
TpS
(9)
AR
(5)
c
dc
D
b
ab ab
a
a
FG J
4
2
0
6
4
2
0
6
4
2
0
IndividualsSingle-species-
dominated
ecosystems
Multispecies-
dominated
ecosystems
FIGURE 6.4
Box plot of light use efciency (LUE) values comparing values between organizational lev-
els and land cover types. The graphic explains the minimum, maximum, rst quartile (25%),
median, mean, and third quartile (75%) of LUE values. The mean is displayed with a +, and a
black line corresponds to the median. Letters indicate signicantly different groups (Kruskal–
Wallis test, p < 0.05). Lowercase letters indicate signicantly different groups within each
organizational level and between land cover types. Uppercase letters indicate signicantly
different groups between organizational levels and per each land cover type. Total LUE values
= 185. Archibold’s land cover type classication: PHM = polar and high mountain tundra, CF
= coniferous forests, TW = terrestrial wetlands, Cr=crops, TFE = temperate forest ecosystems,
ME = Mediterranean ecosystems, TG = temperate grasslands, TpF = tropical forests, TpS =
tropical savannas; and AR = arid regions.
K14591_C006.indd 113 28/08/13 9:11 PM
114 Earth Observation of Ecosystem Services
LUE (g C/MJ APAR)LUE (g C/MJ APAR)LUE (g C/MJ APAR)
6
4
2
0
6
4
2
0
6
4
2
0
Annual
(85)
Seasonal
(23)
Daily
(27)
IndividualsSingle-species-
dominated ecosystems
Multispecies-
dominated ecosystems
BD
a
A
b
ddd
c
C
AD
a
FIGURE 6.5
Box plot of temporal variation of average light use efciency (LUE) values in each organi-
zational level and the estimate periods. The graphic explains the minimum, maximum, rst
quartile (25%), median, mean, and third quartile (75%) of LUE values. The mean is displayed
with a +, and a black line corresponds to the median. Different letters indicate signicant
differences (Kruskal–Wallis test, p < 0.05). Lower case letters indicate signicantly different
groups between time interval of estimation and within each organizational level. Capital let-
ters indicate signicantly different groups between organizational levels and the time interval
of estimation. Total LUE values = 185. Archibold’s land cover type classication: PHM = polar
and high mountain tundra, CF = coniferous forests, TW = terrestrial wetlands, Cr = crops,
TFE= temperate forest ecosystems, ME = Mediterranean ecosystems, TG = temperate grass-
lands, TpF = tropical forests, TpS = tropical savannas, and AR = arid regions.
K14591_C006.indd 114 28/08/13 9:11 PM
115Missing Gaps in the Estimation of the Carbon Gains Service
6.4 Conclusions
Monteith’s model, based on LUE and remotely sensed estimates of fPAR,
constitutes the most widely used approach for mapping the terrestrial carbon
cycle (Jenkins etal. 2007; Pereira etal. 2007), though it is not free of uncertain-
ties. The inherent spatiotemporal variability found among different meth-
odologies may explain the variability of LUE estimates found in this study.
The time interval of the estimates and the level of organization are two clear
sources of such variation. In such a way, annual estimates of NPP at regional
scale should not be used for LUE estimation at individual level and derive
for short-term (e.g., days) measurements. The variability of LUE estimates
related to environmental and physiological factors (such as leaf form, ribu-
lose diphosphate carboxylase content, temperature, and/or moisture) (Ito and
Oikawa 2007; Tong etal. 2008) can result in large errors if these values are
extrapolated to global or regional scales (Nouvellon etal 2000; Piñeiro etal.
2006). Our results indicated that high temporal variation in LUE estimates
at the individual and multispecies-dominated ecosystems levels across land
cover types (see also Grace etal. 2007; Cook etal. 2008; Hilkeretal. 2008) do
not account for regional and global NPP estimates, which typically apply a
constant LUE value (Drolet etal. 2008; Maselli etal. 2009).
Acknowledgments
The authors gratefully acknowledge the staff of the Andalusian Government’s
Department of the Environment for providing the facilities required to obtain
the necessary assistance. They also thank Gervasio Piñeiro, Dolores Arocena,
Carlos Di Bella, Piedad Cristiano, and two anonymous reviewers for their usefu l
comments. Financial support was provided by the ERDF (FEDER), Andalusian
Regional Government (Junta de Andalucía GLOCHARID & SEGALERT
Projects, P09–RNM-5048), and the Ministry of Science and Innovation (Project
CGL2010-22314). Support for A. J. C. was also provided by the Centro Andaluz
para la Evaluación y Seguimiento del Cambio Global (CAESCG) and the
Oklahoma Biological Survey (OBS) at the University of Oklahoma.
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