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Use of a SPAD-502 meter to measure leaf chlorophyll concentration in

Arabidopsis thaliana

Article in Photosynthesis Research · February 2011

DOI: 10.1007/s11120-010-9606-0 · Source: PubMed

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Use of a SPAD-502 meter to measure leaf chlorophyll

concentration in Arabidopsis thaliana

Qihua Ling, Weihua Huang1, and Paul Jarvis*

Department of Biology, University of Leicester, University Road, Leicester LE1 7RH, United
Kingdom.

1
Present address: Shanghai Institute of Plant Physiology and Ecology, Shanghai Institutes for
Biological Sciences, Chinese Academy of Sciences, 300 Fenglin Road, Shanghai 200032,
China.

* To whom correspondence should be addressed e-mail: rpj3@le.ac.uk

tel.: +44 116 223 1296
fax: +44 116 252 3330
 Ling et al., page 2 of 13

Abstract

The SPAD-502 meter is a hand-held device that is widely used for the rapid, accurate and
non-destructive measurement of leaf chlorophyll concentrations. It has been employed
extensively in both research and agricultural applications, with a range of different plant
species. However, its utility has not been fully exploited in relation to the most intensively
studied model organism for plant science research, Arabidopsis thaliana. Measurements with
the SPAD-502 meter produce relative SPAD meter values that are proportional to the amount
of chlorophyll present in the leaf. In order to convert these values into absolute units of
chlorophyll concentration, calibration curves must be derived and utilized. Here, we present
calibration equations for Arabidopsis that can be used to convert SPAD values into total
chlorophyll per unit leaf area (nmol/cm2; R2 = 0.9960) or per unit fresh weight of leaf tissue
(nmol/mg; R2 = 0.9809). These relationships were derived using a series of Arabidopsis
chloroplast biogenesis mutants that exhibit chlorophyll deficiencies of varying severity, and
were verified by the subsequent analysis of senescent or light-stressed leaves. Our results
revealed that the converted SPAD values differ from photometric measurements of solvent-
extracted chlorophyll by just ~6% on average.

Keywords

Arabidopsis; Calibration curve; Chlorophyll content; Chloroplast development; SPAD meter

 Ling et al., page 3 of 13

Introduction

Leaf chlorophyll concentration is an important parameter that is frequently measured as an

indicator of chloroplast development, photosynthetic capacity, leaf nitrogen content, or
general plant health. In the laboratory, it is commonly determined photometrically following
extraction of the pigments using an organic solvent, such as acetone or dimethyl formamide
(Arnon 1949; Porra et al. 1989). While this method is well-established and accurate, it is time-
consuming, destructive (the leaf material must be excised from the plant, and is lost), and
necessitates the use of toxic or flammable chemicals. The SPAD-502 meter (Konica-Minolta,
Japan) provides an alternative method for the measurement of relative leaf chlorophyll levels
that overcomes these disadvantages. It is an inexpensive, hand-held device based on two
light-emitting diodes and a silicon photodiode receptor, that measures leaf transmittance in the
red (650 nm; the measuring wavelength) and infrared (940 nm; a reference wavelength used
to adjust for non-specific differences between samples) regions of the electromagnetic
spectrum. These transmittance values are used by the device to derive a relative SPAD meter
value (typically between 0.0 and 50.0) that is proportional to amount of chlorophyll in the
sample (Uddling et al. 2007). The meter has been used extensively in both research and
agricultural settings, and there are many publications in the scientific literature that describe
its use.
In order to convert relative SPAD meter values into units of absolute chlorophyll
concentration, it is necessary to employ a calibration curve (Markwell et al. 1995). The
relationship between SPAD values and chlorophyll concentration has been investigated in a
variety of different species, and has been found to display considerable interspecific variation
(Castelli et al. 1996; Uddling et al. 2007). This variability is presumed to be due to structural
differences between the leaves of different species, causing different light reflection or
scattering effects. Thus, in order for the SPAD-502 meter to be used to record real chlorophyll
concentration values with reasonable accuracy, a calibration equation must be derived
specifically for the particular species of interest (Richardson et al. 2002). Arabidopsis
thaliana has been widely adopted across the globe as a model organism for studies on plant
development, cellular and molecular biology, and biochemistry. The particular advantages of
this species are well documented, and include its completely-sequenced genome, diminutive
stature, and rapid generation time, as well as the availability of extensive collections of
insertional mutants such that it is possible to quickly identify a knockout for almost any gene
 Ling et al., page 4 of 13

of interest. As a result, a considerable proportion of the chloroplast and plastid research that is
undertaken today is conducted using the Arabidopsis model system (Leister 2003; López-Juez
2007; Sakamoto et al. 2008). Nonetheless, only a very small number of Arabidopsis studies
have employed the SPAD-502 meter, and these have either presented the data in unconverted
arbitrary units (Stettler et al. 2009) or utilized a crude scaling factor to estimate actual
chlorophyll content (Takami et al. 2010). To enable the advantages of the SPAD meter to be
exploited more fully in the field of Arabidopsis research, we have derived specific calibration
relationships that can be used to convert SPAD values accurately into absolute chlorophyll
concentration values, on either a per-leaf-area or a per-fresh-weight basis.

Materials and methods

Plant material and growth conditions

All plants used in this study were Arabidopsis thaliana of the Columbia-0 ecotype. The
+/tic110-1, hsp93-V-1, tic40-4, ppi1-1 and hsp93-V-1 hsp93-III-1 (hsp93-V/III) mutants
employed to derive the calibration relationships have all been described previously (Jarvis et
al. 1998; Kovacheva et al. 2005; Kovacheva et al. 2007). Similarly, the pph-1 and pao1
mutants have been presented in earlier reports (Pruzinska et al. 2005; Schelbert et al. 2009).
Plants were germinated on MS medium and allowed to grow for 10 days for prior to
transferral to soil, or sown on soil directly. They were grown in ~100 μmol/m2/s white light
under a long-day cycle (16-hours-light/8-hours-dark), as described previously (Aronsson and
Jarvis 2002).

Dark and light-stress treatments

Dark treatments for the induction of senescence were conducted using two different methods,
as described previously (Pruzinska et al. 2005; Schelbert et al. 2009). In the first method (Fig.
2), developmentally-equivalent leaves were detached from a plant and placed on wet filter
paper in sealed Petri dishes. The dishes were then placed in a dark container within a growth
cabinet for six days. In the second method (Fig. 3), the leaves were carefully wrapped in
 Ling et al., page 5 of 13

aluminium foil whilst still attached to the plant, and then left under standard growth
conditions for 4-6 days.
Light stress treatments involved exposure of 21-day-old plants grown under standard
conditions to high-intensity white light (~2000 μmol/m2/s), delivered through a chilled-water
heat sink, for three hours per day for seven days. After the final period of high-light treatment,
the plants were returned to standard growth conditions for 24 hours prior to conducting the
measurements.

Chlorophyll measurements and data analysis

Absolute chlorophyll concentration measurements were conducted using a spectrophotometer

and dimethyl formamide extracts of leaf tissue, as described previously (Constan et al. 2004;
Porra et al. 1989). Leaf tissue for these measurements was harvested using a circular punch
(cork borer) that yields 0.5 cm diameter leaf discs that are 0.19635 cm2 in area; excised discs
were also weighed, enabling the chlorophyll data to be expressed in relation to fresh weight as
well as leaf area. Each solvent extract contained several leaf discs from multiple different
plants per genotype or treatment. SPAD values were recorded using exactly the same leaves
from the same plants, prior to sampling, using a SPAD-502 meter (Konica-Minolta, Japan).
Twelve independent SPAD measurements were made per genotype or treatment, using several
different plants.
Relationships between the two datasets were analysed using both SigmaPlot (SPSS
Science Inc.) and Excel (Microsoft) software. Several models were tested and the two
packages gave identical results, indicating that second-order polynomial functions provide the
best fit.

Results and discussion

In order to derive useful relationships between SPAD meter readings and chlorophyll content,
it is necessary to study plants that have a variety of different pigmentation levels. To this end,
we exploited a range of well-characterized chloroplast biogenesis mutants that are defective in
the import of proteins into plastids (Jarvis 2008). The selected mutant genotypes (+/tic110,
 Ling et al., page 6 of 13

hsp93-V, tic40, ppi1 and hsp93-V/III) exhibit varying degrees of chlorosis due to variations in
the importance of the affected genes (Fig. 1a), and were analysed alongside wild-type plants
of two different ages. All of the plants were used contemporaneously to make both SPAD
meter readings and photometric measurements using pigment extracts. By plotting these two
datasets against each other, we assessed possible relationships between them. While linear and
exponential relationships between SPAD values and chlorophyll concentrations have
previously been proposed (Uddling et al. 2007), we observed a much stronger fit using
second-order polynomial functions (Fig. 1, b and c); R2 values for linear and exponential
relationships were, respectively, 0.979 and 0.973 for the data in b, and 0.959 and 0.972 for the
data in c, and thus significantly lower than those shown in Fig. 1 for the selected polynomial
functions. Others have similarly reported that such polynomial functions best describe the
relationship between SPAD values and chlorophyll concentration (Hawkins et al. 2009;
Markwell et al. 1995).
To test the utility of the calibration equations presented in Fig. 1, we studied plants
exhibiting dark-induced senescence. Studying such environmentally induced chlorosis
enabled us to assess whether the aforementioned relationships (derived based on an analysis
of genetically induced chlorosis) would remain valid under different conditions. Two different
methods of dark treatment for the induction of senescence have been described in the
literature: the incubation of detached leaves in a dark container, and the covering of attached
leaves with aluminium foil (Pruzinska et al. 2005; Schelbert et al. 2009). We elected to
employ both of these methods, starting with the former. In addition to wild type, we
incorporated two well-characterized, senescence-defective mutants in our analysis, as
additional controls. These were the pheophytinase (PPH) mutant, pph, and the pheide a
oxygenase (PAO) mutant, pao1, both of which are deficient in an important step of
chlorophyll breakdown and, therefore, express a “stay-green” phenotype under conditions that
normally induce senescence (Pruzinska et al. 2005; Schelbert et al. 2009).
After six days in the dark, detached wild-type leaves were pale-yellow in appearance,
whereas pph and pao1 mutant leaves retained considerable amounts of green pigmentation
(Fig. 2a). While the pph leaves were uniformly green, pao1 leaves had a patchy appearance
with both green and yellow sectors. This phenotypic difference, although not specifically
reported previously, presumably reflects the fact that pheide a and red chlorophyll catabolites
accumulate in pao1, but not in pph, which can cause a lesion-mimic or cell death phenotype
in the former mutant in the light (Schelbert et al. 2009). As a result of its patchy appearance,
pao1 was deemed unsuitable for the purposes of our experiment (as it would have been
 Ling et al., page 7 of 13

difficult to obtain consistent measurements), and so we focused only on the wild type and pph.
Dark-treated and control (untreated) leaves were used to make both SPAD meter readings and
photometric measurements using pigment extracts. The SPAD data were converted to
chlorophyll values employing the calibration equations displayed in Fig. 1, and then
compared with photometrically determined data. As shown in Fig. 2 (b and c), the two
datasets were very similar, indicating that the derived conversion formulae are suitable for the
measurement of chlorophyll under different conditions.
In relation to chlorophyll a/b ratios, it should be noted that the meter’s measuring light
has a peak wavelength of 650 nm, which lies approximately midway between the absorption
maxima of chlorophylls a and b, and in fact is the point of intersection of the absorption
spectra of the two pigments (Arnon 1949; Porra et al. 1989). Thus, while caution is advised,
measurements with the SPAD meter are unlikely to be strongly distorted by a/b ratio changes.
Indeed, the effect of the pph mutation is such that, under dark-treatment conditions, the
chlorophyll a/b ratio is dramatically reduced in the mutant relative to wild type (data not
shown; Schelbert et al. 2009). The fact that SPAD readings taken using dark-treated pph
plants were similar to corresponding photometric measurements (Fig. 2, b and c) supports the
notion that the meter is not adversely influenced by deviations in the chlorophyll a/b ratio.
Using the second method for the induction of senescence, we observed that most
leaves covered for six days would dry out and die. The covering period was therefore reduced
to four days, and under these conditions we observed the expected chlorosis in wild type, and
a “stay-green” phenotype in pph (Fig. 3a). However, pao1 leaves were not viable under these
conditions either, and, as with the detached leaves (Fig. 2), this difference from pph may be
due the accumulation of harmful chlorophyll catabolites in the pao1 mutant (Pruzinska et al.
2005; Schelbert et al. 2009). Importantly, analysis of chlorophyll levels in these leaves using
the two different methods again revealed closely correlated datasets (Fig. 3b), supporting the
utility of the SPAD conversion formulae.
In a final experiment to corroborate the usefulness of the calibration equations, we
analysed light-stressed plants simultaneously using both chlorophyll quantification methods.
Prolonged exposure to high light intensities of a plant not acclimated to such conditions leads
to significant damage of the photosynthetic apparatus and a concomitant depletion of
photosynthetic pigments (Aronsson et al. 2006; Powles 1984). As expected, the light-stressed
plants appeared paler than control plants, and somewhat stunted (Fig. 4a). Comparison of the
converted SPAD data with those derived photometrically from extracted pigment confirmed
the suitability of the formulae for assaying chlorophyll content differences induced by another,
 Ling et al., page 8 of 13

different set of conditions (Fig. 4b). Considering all of the data shown in Figs. 2, 3 and 4, it is
apparent that the values determined using the SPAD meter differ from the conventionally
acquired data by ~6%, on average, which for most purposes should be an acceptable margin
of error. It is noteworthy that there is a strong linear relationship between readings of the
SPAD-502 meter, employed here, and another popular chlorophyll meter, the N-Tester (Yara,
Norway; formerly Hydro) (Levey and Wingler 2005; Uddling et al. 2007). Thus, the
conversion formulae derived here may also be useful in conjunction with data from that
device.
It should be noted that SPAD meter readings can be influenced by changing growth
conditions that, for example, may lead to a redistribution of chloroplasts within mesophyll
cells (Naus et al. 2010). For this reason, it is advisable to always include an internal control in
each experiment, as a point of reference, and to be cautious about making comparisons
between experiments conducted on different occasions. Nonetheless, provided that the plants
to be measured are grown under the commonly-used and very standard, controlled conditions
employed here, the results should be reproducible. Another consideration that should be borne
in mind is the size of the meter’s measuring area, which is 2 mm × 3 mm. While this
compactness is an advantage in that it may accommodate all but the smallest leaves of rosette-
stage plants, it may be disadvantageous in relation to leaves that have an excessively patchy
or reticulate appearance (e.g., pao1; Fig. 2a). However, this problem may be at least partially
overcome by taking multiple measurements using different areas of the same leaf.

Conclusion

The formulae presented here are suitable for converting SPAD readings into values of
chlorophyll concentration in Arabidopsis leaves, and the converted values correlate closely
with direct photometric measurements of extracted chlorophyll. Use of the SPAD-502 meter
is a very quick and simple procedure, requiring no toxic or flammable solvents. Moreover, as
the procedure is non-destructive, this approach has significant advantages in relation to
experiments that require the recovery of biological material from the leaves (e.g., nucleic
acids or protein), the analysis of particularly precious plants that can neither be sacrificed nor
damaged, and studies where it is desirable to monitor dynamic changes of chlorophyll content
in the same individual throughout development or in responses to changing conditions.
 Ling et al., page 9 of 13

Acknowledgements

This work was supported by a Royal Society International Incoming Fellowship (to WH), the
Royal Society Rosenheim Research Fellowship (to PJ), and by Biotechnology and Biological
Sciences Research Council (BBSRC) grants BB/D016541/1, BB/F020325/1 and
BB/H008039/1 (to PJ).
 Ling et al., page 10 of 13

Figure legends

Fig. 1. Use of chloroplast biogenesis mutants to derive calibration relationships between

SPAD meter readings and photometric chlorophyll measurements.
(a) Arabidopsis plants of the indicated genotypes were grown side-by-side under identical
conditions for either 35 days (wild type and all mutants) or 45 days (wild type only), and then
typical individuals were selected for photography.
(b, c) Plants similar to those shown in (a) were used to make SPAD meter readings, and to
make direct measurements of absolute chlorophyll concentration by the photometric analysis
of solvent extracts. The SPAD values are plotted along the x-axis, and the corresponding
chlorophyll concentration values are plotted along the y-axis (nmol chlorophyll a+b per cm2
leaf area in [b]; nmol chlorophyll a+b per mg leaf fresh weight in [c]). The presented data are
means (±SD) derived from 12 independent SPAD meter readings done on up to 12 different
plants, or from three independent solvent extracts (one less for tic40), each one containing
four leaf discs from up to four different plants. Second order polynomial functions were found
to fit the data most closely; the relevant curves and equations, and corresponding R2 values,
are shown in each panel.

Fig. 2. Analysis of dark-induced senescence in detached leaves for the validation of the
calibration equations.
(a) Developmentally equivalent leaves of 35-day-old wild-type and “stay-green” mutant (pph
and pao1) plants were detached, incubated on wet filter paper in a dark container for six days,
and then photographed.
(b, c) Leaves similar to those shown in (a) were used to make both SPAD meter readings and
photometric measurements of chlorophyll concentration using extracted pigments. The SPAD
readings were converted to chlorophyll concentration values using the relationships presented
in Fig. 1, and then the two datasets were plotted side-by-side for comparison purposes. Values
shown are means (±SD) derived from 12 independent SPAD meter readings done on six
different plants, or from three independent solvent extracts, each one containing four leaf
discs from two different plants.
 Ling et al., page 11 of 13

Fig. 3. Analysis of dark-induced senescence in attached leaves for the validation of the
calibration equations.
(a) Developmentally equivalent leaves of 35-day-old wild-type and “stay-green” mutant (pph
and pao1) plants were covered with aluminium foil (whilst still attached to the plant), and
then left for four days under standard growth conditions prior to detachment and photography.
(b) Leaves similar to those shown in (a) were used to make both SPAD meter readings and
photometric measurements of chlorophyll concentration using extracted pigments; the SPAD
readings were converted to chlorophyll concentration values using the relationships in Fig. 1
prior to analysis. Values shown are means (±SD) derived from 12 independent SPAD meter
readings done on six different plants, or from three independent solvent extracts, each one
containing four leaf discs from two different plants.

Fig. 4. Analysis of light-stressed plants for the validation of the calibration equations.
(a) Wild-type Arabidopsis plants were grown under standard conditions for 21 days, and then
subjected to periods of high-light stress (2000 μmol/m2/s) over a seven day period. Typical
stressed and control (untreated) plants are shown.
(b) Developmentally equivalent leaves from plants similar to those shown in (a) were used to
make both SPAD meter readings and photometric measurements of chlorophyll concentration
using extracted pigments; the SPAD readings were converted to chlorophyll concentration
values using the relationships in Fig. 1 prior to analysis. Values shown are means (±SD)
derived from 12 independent SPAD meter readings done on three different plants, or from
three independent solvent extracts, each one containing four leaf discs from a single plant.
 Ling et al., page 12 of 13

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Figure 1
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Figure 2
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Figure 3
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Figure 4
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Erratum to: Use of a SPAD-502 meter to measure leaf
chlorophyll concentration in Arabidopsis thaliana

Photosynth. Res. (2011) 107:209–214

DOI 10.1007/s11120-010-9606-0

Qihua Ling, Weihua Huang1, and Paul Jarvis*

Department of Biology, University of Leicester, University Road, Leicester LE1 7RH, United
Kingdom.

1
Present address: Institute of Plant Physiology and Ecology, Shanghai Institutes for
Biological Sciences, Chinese Academy of Sciences, 300 Fenglin Road, Shanghai 200032,
China.

* To whom correspondence should be addressed e-mail: rpj3@le.ac.uk

tel.: +44 116 223 1296
fax: +44 116 252 3330
 Ling et al., page 2 of 2

Due to the omission of a scaling factor of 4 from the chlorophyll per leaf area calculations, all
values with units of nmol/cm2 were fourfold higher than they should have been. This affected
the y-axis values in Figs. 1b, 2b, 3b (left side) and 4b (left side); the maximal y-axis values
should be 30, 25, 15 and 35, respectively. Most importantly, the equation in Fig. 1b should be:

y = 0.0105x2 + 0.4119x + 0.3810.

None of the chlorophyll per fresh weight data are affected by this erratum, nor is the running
text influenced in any way. All R2 values are unaffected.

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