PHOTOSYNTHETICA 48 (4): 596-609, 2010

Modelling photosynthetic photon flux density and maximum potential

gross photosynthesis
R.J. RITCHIE
Faculty of Technology and Environment, Prince Songkla University Phuket, Thailand 83120

Abstract
Irradiance data software developed by the NREL Solar Radiation Laboratory (Simple Model of Atmospheric Radiative
Transfer of Sunshine, SMARTS) has been used for modelling photosynthesis. Spectra and total irradiance were expressed
in terms of quanta [mol m2 s1, photosynthetic photon flux density, PPFD (400700 nm)]. Using the SMARTS software
it is possible to (1) calculate the solar spectrum for a planar surface for any given solar elevation angle, allowing for the
attenuating effects of the atmosphere on extraterrestrial irradiance at each wavelength in the 400700 nm range and for
the thickness of atmosphere the light must pass through during the course of a day, (2) calculate PPFD vs. solar time for
any latitude and date and (3) estimate total daily irradiance for any latitude and date and hence calculate the total photon
irradiance for a whole year or for a growing season. Models of photosynthetic activity vs. PPFD are discussed. Gross
photosynthesis (Pg) vs. photosynthetic photon flux density (PPFD) (Pg vs. I) characteristics of single leaves compared to
that of a canopy of leaves are different. It is shown that that the optimum irradiance for a leaf (Iopt) is the half-saturation
irradiance for a battery of leaves in series. A C3 plant, with leaves having an optimum photosynthetic rate at 700 mol
m2 s1 PPFD, was used as a realistic worked example. The model gives good estimates of gross photosynthesis (Pg) for
a given date and latitude. Seasonal and annual estimates of Pg can be made. Taking cloudiness into account, the model
predicts maximum Pg rates of about 10 g(C) m2 d1, which is close to the maximum reported Pg experimental
measurements.
Additional keywords: global models; gross photosynthesis; irradiance; light saturation curves; modelling; photoinhibition;
photosynthetically active radiation; photosynthetic photon flux density; primary productivity.

Introduction
For most physical applications and in climatology solar
energy is given in W m2; however, in photosynthetic
applications, because of the quantum nature of the light
reactions of photosynthesis, biologists are more interested
in irradiance presented as photosynthetically active
radiation (PAR) in units of mol m2 s1 (McCree 1973).
The unambiguous name for PAR expressed in quantum
terms is photosynthetic photon flux density (PPFD).
For a given wavelength of monochromatic light
irradiance in terms of mol m2 s1 and W m2 can be
interconverted using the Planck equation:

P =

N A hnc

or solving for n,
n =

P
N A hc

(1)

where P is the electromagnetic power delivered,

expressed on a surface area basis (W m2), NA is the
Avogadro constant (6.0221367 1023 mol1), n is the

Received 25 March 2010, accepted 15 October 2010.

Present address: School of Biological Sciences A-08, The University of Sydney, NSW 2006, Australia.
Phone: +61 9351 4475, fax: +61 2 9351 4119, e mail: rrit3143@usyd.edu.au
Abbreviations: Chl chlorophyll; ETI extraterrestrial irradiance; ETR electron transport rate; I irradiance PPFD; Id daily
irradiance PPFD, Pg gross photosynthesis, PAR photosynthetically active radiation, PPFD photosynthetic photon flux density,
PS photosystem; SMARTS Simple Model of Atmospheric Radiative Transfer of Sunshine; p photosynthetic efficiency.
Acknowledgements: The author wishes to thank Dr John W. Runcie (University of Sydney), Emeritus Prof Tony Larkum and Mr
Mark Curran (ret.) (University of Sydney) for their interest in this study and helpful comments on the paper. I especially wish to thank
Dr Christian Gueymard for help in using the SMARTS software (NREL and Solar Consulting Services http://
www.SolarConsultingServices.com). This project was funded by an Endeavour Executive Award to the author (No 1324_2009, Dept
of Education and Training, Commonwealth of Australia).

596

PPFD AND PHOTOSYNTHESIS

number of moles of quanta per second on a surface area

basis (mol m2 s1), h is Plancks constant (6.6260755
1034 J s), c is the speed of light in vacuum (2.99792458
108 m s1), is the wavelength (m).
Unless the wavelength is specified it is not possible to
interconvert PAR irradiance expressed in W m2 and
PPFD in mol m2 s1. One mole of quanta is sometimes
called one einstein but is not an official SI unit. For some
types of light source there are approximate values for
interconversion of PAR (400700 nm) expressed
in various units (Gensler 1984) but care needs to be taken
in their use.
Very reliable extraterrestrial spectra of the sun have
now been published (Gueymard et al. 2002, Reference
Solar Spectral Irradiance 2003). Dr. C. Gueymard
developed the SMARTS software (SMARTS 2009,
http://www.nrel.gov/rrdc/smarts/) to model solar radiation reaching the earths surface (Gueymard 1995,
Gueymard 2001, SMARTS 2009). The NREL Solar
Radiation Laboratory (http://www.nrel.gov/) hosts the
SMARTS webpage and has developed the Excel interface.
Four properties of photosynthesis are often misunderstood when attempts are made to estimate potential
primary productivity from irradiance data. These are; the
quantum nature of the photosynthetic mechanism
imposes a strict upper limit on carbon fixation rates, only

part of the PPFD spectrum (400700 nm) is actually used

in photosynthesis and the photosynthetic mechanism, like
most biochemical processes, is not only saturable but in
addition substantial photoinhibition can occur at high
irradiances.
In the present study I have developed a minimal
model of photosynthesis based on accurate modelling of
irradiance in quantum units. Using the SMARTS software
(Gueymard 1995, Gueymard 2001, SMARTS 2008) and
the NREL: Solar Position Calculators: SOLPOS it is
possible to calculate irradiance at any latitude and solar
elevation angle (corrected for refraction) over the period
of a day for a given date. Numerical integration methods
can then be used to calculate daily and yearly irradiances
for a given latitude. Given a realistic model of photosynthesis that takes photoinhibition into account and
allows for only part of the photosynthetically active
radiation spectrum (400700 nm) being actually useable
for photosynthesis, it is possible to make realistic
estimates of primary productivity for a given latitude and
time of year. The models developed here are deliberately
as simple and general as possible but take advantage of
better irradiance data (Gueymard 1995, Gueymard 2001,
NREL Solar Radiation Laboratory 2008, SMARTS 2009)
and a better model of photosynthesis vs. irradiance.

Theory: development of a model of gross photosynthesis

Fig. 1 shows that plots of PPFD vs. wavelength, even for
the suns extraterrestrial spectrum, are not the smooth
curves expected from a black body with a colour temperature of about 5,770 K. There are substantial
absorption bands arising from the presence of elements in
the chromosphere of the sun, in particular near 450 nm.
The solar spectrum which reaches the ground is
considerably modified by passing through the atmosphere
(plotted in Fig. 1 as total irradiance upon a flat horizontal
surface, global horizontal irradiance total diffuse and
direct sunlight as defined in the SMARTS software
described in the Methods). Furthermore, the atmosphere
does not behave as a simple neutral density filter, for
example there is a strong absorption band at 687698 nm
and there is a general trend for absorption/scattering to be
greatest for blue light and for the degree of absorption to
decrease towards the red end of the spectrum. This effect
is more pronounced for diffuse horizontal irradiance
(skylight, as defined in the SMARTS software) than for
direct sunlight (direct normal irradiance, as would be
measured with a detector pointed directly at the sun) and
global horizontal irradiance.
The overall photosynthetic reaction for C3 photosynthesis (Larkum et al. 2003, Miller 2006, Falkowski
and Raven 2007) is:
12H2O + 6CO2 + 54 photons (hc/)C6H12O6 + 6H2O + 6O2

(2)

or expressed in the simplest ratios,

2H2O + CO2 + 9 photons (hc/)(CH2O)n + H2O + O2

The theoretical efficiency of photosynthesis based

upon the energy in 54 moles of photons with a
wavelength of 680 nm (the red peak of Chl a in vivo)
(Planck equation) and the energy required to synthesise
one mole of glucose (2,803 kJ mol1) is about 29.5%.
This upper limit for photosynthetic efficiency is often
incorrectly thought to show that there is great room for
improvement in the photosynthetic efficiency of plants
growing in sunlight. Some of the reasons why this is an
incorrect inference are:
(1) It is not possible for oxygenic photosynthesis to
function with less than 9 photons per mole of carbon
fixed: even 8 photons of red light will not provide enough
ATP and NADPH+H+ to fix one CO2 molecule using the
Calvin cycle (Larkum et al. 2003, Miller 2006, Falkowski
and Raven 2007).
(2) 9 photons of blue light (Chl a peak 440 nm)
cannot fix any more carbon than 9 photons of red light at
680 nm even though the blue photons have 680/440 or
54.5% more energy. This is why photosynthetic calculations should always use quantum units rather than
radiometric units (W m2).
(3) Not all photons between 400 and 700 nm
wavelengths are equally useable in photosynthesis. The

597

R.J. RITCHIE

light harvesting by photosynthetic pigments and the

quantum efficiency of photons actually absorbed varies
considerably at different wavelengths. That is after all
why leaves appear green in colour. For green algae and
vascular plants containing Chl a and b (Larkum et al.
2003, Miller 2006, Falkowski and Raven 2007) in terms
of the proportion of photons used for photosynthesis, the
efficiency of the use of photons for photosynthesis is high
near the in vivo blue and red absorption peaks of
chlorophyll (Chl) a (440 & 680 nm respectively) but very
low for green light (550 nm). The solar spectrum at wavelengths less than 425 nm is very depleted, particularly at
ground level (Fig. 1) and so far-blue and violet light
would contribute very little to total photosynthesis.
Xanthophylls and carotenoids are accessory photosynthetic pigments, which absorb energy and transfer energy
to the photoreaction centres of photosystems (PS) I and
II. They absorb light at wavelengths from about 450 to
500 nm and efficiently transfer absorbed energy to PSI
and PSII. These pigments provide an extension of the
window of solar radiation useable for photosynthesis.
Based upon classical action spectra of photosynthesis
(photosynthesis vs. wavelength of photons corrected to
allow for the quantum nature of light) (McCree 1972,
Kyewalyanga et al. 1997, Larkum et al. 2003, Miller
2006, Falkowski and Raven 2007), an estimate of light
actually used for photosynthesis would be the sum of
irradiance from 425 to 500 nm plus irradiance from 640
to 700 nm, allowing for the actual quantum efficiency at
different wavelengths. Thus, an optimistic estimate of
photosynthetically useable irradiance (IPUR) is;
500

I PUR 0.668

I d + 0.819

425

700

(3)

640

where the constant 0.668 is the average quantum

efficiency for blue light (425500 nm) and 0.819 is the
average quantum efficiency for red light (640700 nm)
based on polynomial interpolation of the data of McCree
(1972) for a variety of crop plants. For full sunlight on
earth, at the equatorial equinox, the PPFD is about
2,220 mol m2 s1 but only 34.6% of the total irradiance
(767 mol m2 s1) could actually be used in photosynthesis by Chl a/b-type plants (calculated from Eq. 3
for the standard solar spectrum for a flat surface at
equatorial equinox shown in Fig. 1). Action spectra for
organisms containing accessory Chl c of various types
(diatoms, dinoflagellates and other chromophytes and the
zooxanthellae of corals and clams) are essentially similar
to those found for Chl a/btype organisms (green algae
and vascular plants) (Neori et al. 1988, Khl et al. 1995,
Kyewalyanga et al. 1997). In the present study, sun/shade
adaptation of plants under low irradiance conditions to
utilise as much light as possible in the green-yelloworange parts of the spectrum (500640 nm) and the
package effect arising from self-shading effects of
chloroplasts and light-harvesting pigments (Jones 1992)

598

have been neglected because most of the photosynthesis

of a forest canopy occurs in the top layers exposed to
high irradiance. The overall contribution to total photosynthesis of heavily shaded leaves in a canopy would be
small and so any special adaptations of them to very low
irradiance would have little effect on the total photosynthesis of the canopy. The finding that differences in
the package effect were not large in tropical forest sun
and shade plants (Lee et al. 1990), is also reassuring.
Higher plants and green algae (Hodges and Barber
1983, Finazzi et al. 2002, Iwai et al. 2010) are able to use
State I/State II transitions to adjust their photosynthetic
mechanism to ambient conditions without the synthesis
of new photosystems and antennae complexes. Thus to
some extent they are able to adjust their photosynthetic
action spectra to continuously fit their ambient light
regime.
Cyanobacteria (blue-green algae), rhodophytes (red
algae), glaucocystophytes and cryptophytes have
phycobiliproteins which harvest green and yellow light,
at least under low-light conditions (Subramaniam et al.
1999). Such photoautotrophs have much more dramatic
rapid photoadaptation effects than in vascular plants and
green algae. Eq. 3 cannot be easily adapted to apply to
such algae by simply inserting a term to take light
absorbed by phycobiliproteins into account. It is difficult
to estimate how much of the PPFD window is usefully
absorbed by phycobiliproteins because cells containing
these accessory pigments are able to photo-adapt very
quickly by coupling or decoupling their phycobilisomes
from their photosystems (Subramaniam et al. 1999).
Thus, in low irradiances phycobilin-containing algae
would be able to use most absorbed light in the PPFD
spectrum (400700 nm) with high quantum efficiency but
under high light conditions the phycobiliproteins would
be decoupled and the cells would have an action spectrum
similar to that of green algae and vascular plants (Eq. 3)
(Ploug et al. 1993). Thus although phycobilin pigmentation of cyanobacteria confers an advantage under very
low irradiances (Raven et al. 2000, Falkowski and Raven,
2007), in an open-air algal production pond under full
sunlight the phycobilins confer less energetic advantage,
compared to a green alga than usually thought because
under such conditions the phycobilins are largely
decoupled (Ploug et al. 1993, Subramaniam et al. 1999).
Objections to applying Eq. 3 to all oxyphotoautrophs
turn out to be less justifiable than they at first appear to
be. Eq. 3 is not an unreasonable approximation for
modelling total photosynthesis by any community of
oxyphotoautotrophs provided that (1) irradiance is high
and (2) they use Chl a as the primary photochemical
pigment. A forest, a dense crop, a bank of corals, an algal
bloom, a cyanobacterial production pond or an algal mat
should all use no more than about 3040% of incident
PPFD. Eq. 3 does not apply to the exotic cyanobacterium,
Acaryochloris marina, because it uses Chl d (not Chl a)
as its primary electron acceptor and uses light outside the

PPFD AND PHOTOSYNTHESIS

400700 nm range (Gloag et al. 2007, Ritchie 2008).

Eq. 3 would also not apply to oxyphotoautotrophic
bacteria which use the newly-discovered Chl f, which has
a methanol solvent absorption peak at 706 nm (Chen
et al. 2010).
Surface inhibition of photosynthesis is well known to
limnologists and oceanographers: models of primary
productivity of oceanic phytoplankton have included
consideration of the saturation and photoinhibition
properties of the photosynthetic mechanism at high
irradiances for a very long time (Walsby 1997, Miller
2006, Falkowski and Raven 2007, Ritchie 2008). Such
models have been applied to algal mats (Ploug et al.
1993). Nevertheless, light saturation and photoinhibition
properties of the photosynthetic mechanism are often
neglected in models estimating theoretical primary
productivity from irradiance data (Posada et al. 2009).
Asymptotically saturating curves (e.g., MichaelisMenten, exponential saturation, non-rectangular hyperbolae and Tanh) can be used for modelling the response
of photosynthesis to irradiance but such models would
appear to only apply to irradiances below where
photoinhibition sets in (Friend 2001, Larkum et al. 2003,
Miller 2006, Falkowski and Raven 2007, Myeni et al.
2007). The waiting-in-line equation (y = x ex) has been
found to be a very good model for photosynthesis for
suboptimal, optimal and supraoptimal irradiances
for photosynthetic materials that can be considered as
a surface; for example leaves, algal mats, algae filtered
onto glass fibre disks and algal suspensions with a short
light-path. Two forms suitable for modelling photosynthesis (Gloag et al. 2007, Ritchie 2008, Ritchie and
Bunthawin 2010) are,
Pg = Pmax k w Ie

1- k w I

(4a)

or in a form more easily handled in non-linear curve

fitting procedures,
Pg = Pmax .

I
Iopt

1 - I/I opt

(4b)

where Pg is gross photosynthesis measured as electron

transport rate (ETR), O2 evolution or CO2 uptake, Pmax is
the maximum gross photosynthesis, kw is the waiting-inline scaling constant for the PPFD axis, I is the irradiance
(mol m2 s1 PPFD), and Iopt is the irradiance at which
maximum photosynthesis takes place (optimum
irradiance). For the waiting-in-line equation it can be
shown that Pmax occurs when I = 1/kw.
The maximum photosynthetic efficiency (p) is the
initial slope of the curve at I = 0 (p = Pmax e kw). At very
low light intensities photosynthesis is directly proportional to irradiance. The half-maximum photosynthesis
(Phalf-max) occurs at 0.23120 Iopt and photosynthesis is
inhibited by 50% at 2.6734 Iopt (Ritchie 2008).

Eqs. 4a,b apply to a photosynthetic surface that is

opaque. In reality, photosynthesis is carried out by layers
of photosynthetic cells, for example the leaf canopy of
forests, grasslands and crops, algal mats, and water
columns containing phytoplankton. In a community of
photosynthetic cells most cells are at least partially
shaded by cells above them, affording partial protection
from the photoinhibitory effects of excessive irradiance.
Integrated forms of Eqs. 4a or 4b are therefore appropriate for estimating photosynthesis by a uniform but
multi-layered community of photosynthetic units.
The amount of photosynthetically useable radiation
(IPUR) at a given depth (x) inside a translucent
photosynthetic material should approximately obey
Lamberts law (Friend 2001, Suila et al. 2004);
I

= I e

kix

(5)

where x is the depth inside the photosynthetic material, ki

is the attenuation constant and I0 is the PPFD at the
surface of the photosynthetic entity.
For modelling purposes some broad generalisations
need to be made about translucent photosynthetic
materials (leaves of a canopy, algal mat or column of
phytoplankton suspension). Generally the compensation
point (where gross photosynthesis just balances
respiration and hence net photosynthesis is zero) is at
about 0.5% of full sunlight PPFD (e-kix 0.005) and so
the compensation point is at about 0.005 2 224 =
11.1 mol m2 s1 (PPFD) (Raven et al. 2000). A compensation point between 4 and 20 mol m2 s1 is a good
generalisation over a wide range of photoautotrophs,
except those adapted to very low irradiances which never
experience full sunlight (Raven et al. 2000, Larkum et al.
2003, Falkowski and Raven 2007). The value of ki will
vary depending on the actual battery of photosynthetic
units. Representative species in the Panamanian rainforest
have a leaf area index of about 4 to 5 (Posada et al.
2009). On a broader scale, the leaf area index of the
Amazonian rainforest averages 4.7 (Myeni et al. 2007)
and so the ki for such a closed canopy forest would be
1.127 calculated from Eq. 5 for 99.5% attenuation of light
at ground level
It can be shown that taking Eq. 5, substituting into
Eq. 4b and integrating over the depth (x) of the
translucent photosynthetic material it is possible to
estimate the total photosynthesis of a layer of
photosynthetic cells that utilises all the useable light
reaching the depth x in the photosynthetic material.

P =
g

Pmax e I o e k i x /I opt
I /I
(e
e o opt )
ki

(6a)

Eq. 6a is cumbersome and a more general simplified

form is needed here. If we assume that a translucent layer
of photosynthetic cells is thick enough to use virtually all
of the useable light and hence e-kix 0 and so

599

R.J. RITCHIE

eI0e-kix/Iopt 1, a good approximation is a simple

exponential saturation curve (Eq. 6b) which is in form
like the big leaf model used in forest studies (Friend
2001) but saturates very slowly. Note that the optimum
irradiance for a leaf (Iopt from Eq. 4b) is the half
saturation irradiance for a battery of leaves in series. The
waiting-in-line model may be a better descriptor of
photosynthesis vs. PPFD for individual leaves (Eq. 4b)
than the asymptotic rectangular hyperbolae (MichaelisMenten curve) often used in modelling photosynthesis
(Thornley 1998, Friend 2001) but it turns out that a
simple exponential saturation curve does describe
photosynthesis vs. PPFD for a bank of leaves in series.

(7)

where 9 photons are required to fix 1 carbon using C3

photosynthesis (Eq. 2), 0.84 is the average leaf absorptance factor of Bjrkman and Demmig (1987) that is used
in pulse amplitude modulation (PAM) fluorometers as a
standard estimate of the proportion of PPFD irradiance
that is actually absorbed by a photosynthetic surface
(White and Critchley 1999, Gloag et al. 2007, Ritchie
2008), 34.6% of PPFD photons are actually potentially
useable (Eq. 3), an optimum irradiance (Iopt) of 700 mol
m2 s1 PPFD, irradiance (I) is for the given time, date
and latitude as PPFD.
Fig. 2 shows a plot of the Waiting-in-Line function
for a Pmax of 100% and an optimum irradiance (Iopt) of
700 mol m2 s1 PPFD (k = 0.001429) where photosynthesis is expressed on a surface area basis (Eq. 4a) as
well as for a translucent body absorbing and utilising all
useable incident light (Eq. 7). A saturating irradiance of
about 700 mol m2 s1 PPFD would be representative for
most C3 vascular plants grown in full sunlight (Hodges
and Barber 1983, White and Critchley 1999, Ritchie
2008). A photosynthetic surface with a photosynthetic
optimum of 700 mol m2 s1 PPFD would photosynthesise at half the maximum rate in an irradiance of only
161 mol m2 s1 PPFD but would also be inhibited 50%
at 1,871 mol m2 s1 PPFD (Eqs. 4a,b). A multilayered
photosynthetic material (canopy of leaves, algal mat or a
water column containing phytoplankton) which consists
of many layers of cells with P vs. I characteristics
described by Eqs. (4a,b) would follow a simple exponential saturation curve (Eq. 7, Fig. 2). At least theoretically,
a translucent photosynthetic surface, which absorbs all
photosynthetically useable incident light, should exhibit
an asymptotic saturation curve with no photoinhibition
even though thin sections on the upper side will show
photoinhibition at high irradiances.

Use of the SMARTS 2.9.5 software: The SMARTS 2.9.5

software (http://www.nrel.gov/rrdc/smarts/ NREL Solar
Radiation Laboratory, Golden, CO, USA) was used to
calculate PPFD spectra (400700 nm) and total PPFD
over time during daylight and daily total PPFD over the
course of a year at a range of latitudes. The SMARTS295
Users Manual PC and SMARTS295i1.3 manuals were
used as guides for configuring the software. The SMARTS
software allows configurations to be checked for conflicts
and stored as input files as *******_INP.txt. The
software output is stored by the program as
SMARTS295_OUT.txt and SMARTS295_EXT.txt. The
SMARTS295_OUT.txt gives information on the conditions set for each run of the software and a summary of
the output, but limited spectral information. The
SMARTS295_EXT.txt provides complete spectral information for the spectral range specified (but can store no
more than 64 spectra when SMARTS is implemented in

Excel). These output files are overwritten on each run of

the software and so have to be renamed for filing.
Irradiance output terms used: direct normal irradiance
irradiance normal to the solar elevation angle (as would
be measured by a collimated radiometer pointed directly
at the sum), diffuse horizontal irradiancescattered
irradiance from the sky, global horizontal irradiance
total irradiance on a flat horizontal surface. Global
horizontal irradiance is calculated by the SMARTS
software but the calculated value is approximately the
sum of the diffuse horizontal irradiance + sin (solar
elevation angle) direct normal irradiance.
The configurations used for the SMARTS software in
the present study are shown in the Appendix. Default
settings were used if appropriate. The U.S. Standard
Atmosphere is the most commonly used atmospheric
setting (Configuration Card 3). Other latitude-specific
atmospheric settings are available as options in the

Pg

Pmax e
ki

(1 e

I o /I opt

(6b)

Taking the photosynthetic equation (Eq. 2) and an

estimate of useable PPFD (Eq. 3) it is possible to estimate
Pg for a given irradiance regime. Leaves of terrestrial
plants grown in full sunlight, such as clover and peas (C3
plants), typically have a saturating rate of photosynthesis
(Iopt) at about 700 mol m2 s1 PPFD (White and
Critchley 1999, Ritchie 2008). The quantity Pmax e/ki
can be taken as the theoretical maximum yield from
photosynthesis for a given PPFD (Pmax = 0.346/9 mol(C)
mol1(photon) and an Iopt of 700 mol m2 s1.
Substituting into Eq. 6b,
0.84 0.346 I (1 e- I/700 )
=
9
= 0.03229 I ( 1 e I/700 )

P =
g

or,

P = 0.3875 I ( 1 e
g

I/700

Methods

600

PPFD AND PHOTOSYNTHESIS

SMARTS software, e.g. tropical and arctic summer and

arctic winter. As yet there are no standard settings for the
Antarctic. The choice of atmospheric model has a
noticeable effect on the irradiance at some specific
wavelengths (see the spectral region 687698 nm on
Fig. 1) but its effect upon total PPFD (400700 nm) is
less than 0.2%. The default atmospheric CO2 (370 ppmv)
used by SMARTS (Configuration Card 7) is out-of-date:
the estimated 2009 CE CO2 level used in the present
study was 389 ppmv based on data from the NOAA Earth
System Research Laboratory (2009). The SMARTS
default tilt is 37o (the standard tilt for solar panels) and
the azimuth is set to 180o (facing directly south for the
northern hemisphere) (Configuration Card 10a). For
photosynthetic applications the tilt needs to be set to
horizontal (zero tilt) and the azimuth value is not relevant
for a flat horizontal surface. The appropriate spectral
range (minimum 400 nm, maximum 700 nm) needs to be
specified twice: on Configuration Cards 11 and 12.

Geometry for the calculation of atmospheric thickness: For the present study, values for the solar elevation
angle () corrected for atmospheric refraction and relative
atmospheric mass (RAM) were obtained from the NREL
solar radiation laboratory website (NREL solar radiation
laboratory using SOLPOS). Standard settings were used
except that all data was calculated for zero longitude
(Greenwich). The solar elevation angle algorithms used
by the SOLPOS software uses the Standard US atmosphere model to allow for the refractive properties of the
atmosphere. 15-min intervals were chosen as suitable for
the purposes of the present study. Solar angle data can
also be accessed through the SMARTS software
(Configuration Card 17). Differences arising from the
choice of atmosphere models (Configuration Card 3) are
only likely to be significant at low solar elevation angles
where the irradiance on a horizontal flat surface would be
low in any case.

Results
Fig. 1 shows the global horizontal irradiance spectra of
sunlight expressed as mol m2 s1 nm1 from 400 to
700 nm calculated using the SMARTS software. Spectra
are shown for extraterrestrial irradiance (ETI), the
equator, Darwin, NT, Australia (12o28S, 130o50E),
tropic of Cancer, 37oN and 55oN. The appropriate
SMARTS atmosphere option was chosen for each of the
above latitudes (tropical and mid-latitude). Atmospheric
attenuation is more severe towards the blue end of the
PPFD spectrum and so the maximum of the terrestrial
spectrum is moved to longer wavelengths compared to
the extraterrestrial spectrum. The lower the solar angle
the more apparent this red-shift becomes because light
has to pass through a thicker layer of atmosphere. Spectra
were calculated for the March equinox for all latitudes
except for Darwin where the September equinox was
used. The emission spectrum of the sun (ETI), expressed

as quanta (mol m2 s1), is different in appearance to

when it is plotted in terms of energy (W m2). The maxima for solar emission are also different. The maximum
for emitted photons is at 584 nm (9.1 mol m2 s1 nm1).
Comparison of the ETI and spectra at a range of
latitudes for the equinox clearly show that the attenuation
properties of the atmosphere are quite different at
different wavelengths (Fig. 1). The total irradiance for
400 to 700 nm can be summed: at the Equator, the total
PPFD at noon at the equinoxes is 2,209 mol m2 s1
(mean of March and September equinoxes), at 37oN the
total PPFD onto a flat surface at the noon of the spring
equinox is 1,240 mol m2 s1. Similar calculations could
be made at any solar elevation angle using the SMARTS
software (Gueymard 1995, Gueymard 2001, SMARTS
2009).

Fig. 1. Spectral density of PPFD through the

atmosphere onto a planar surface. Solar spectra
expressed in quantum terms for wavelengths from
400 to 700 nm (PPFD). The extraterrestrial
spectrum (ETI) has strong absorption bands from
elements in the chromosphere of the sun. In addition
the atmosphere also has strong absorption bands.
March equinox solar irradiance spectra are shown
for the equator, tropic of Cancer (23o27N), 37oN
and 55oN and the September equinox values are
shown for Darwin (12o28 S). Spectra were calculated using the SMARTS software (SMARTS 2009,
Gueymard 1995, Gueymard 2001). Note the strong
absorption bands near 487 nm and 687698 nm
which are within the blue and red absorption peaks
of chlorophyll a in vivo.

601

R.J. RITCHIE
Table 1. Theoretical primary productivity at various latitudes.
Latitudeo

Solar latitude

Growing season

Daily PPFD Daily carbon

[mol m2 d1] fixation
[g(C) m2 d1]

Total irradiance
Total carbon
in growing season fixation for
[mol m2]
growing season
[g(C) m2]

Tropic of Cancer
23o30N

Summer solstice
Equinox
Winter solstice

All year (365 d)

64.2
52.7
32.3

21.8
17.6
9.90

18,615

6,136

Equator

Solstice
Equinox

All year (365 d)

58.1
52.8

19.8
17.6

20,238

6,823

Darwin
(12o28S)

Summer solstice
Equinox
Winter solstice

All year (365 d)

59.9
56.6
43.4

20.3
19.2
13.9

19,710

6,602

Tropic of Capricorn
23o30S

Summer solstice
Equinox
Winter solstice

All year (365 d)

64.2
52.8
33.3

21.8
17.6
9.90

18,425

6,062

37oN

Summer solstice
Equinox
Winter solstice

7 months
01-Mar to 01-Oct
(214 d)

66.2
44.9
20.2

22.3
14.2
4.98

12,208

4,031

37oS

Summer solstice
Equinox
Winter solstice

7 months
01-Sep to 01-Apr
(212 d)

66.3
45.1
20.2

22.3
14.3
4.98

11,915

3,927

55oN

Summer solstice
Equinox
Winter solstice

5 months
01-May to 01-Oct
(153 d)

64.7
30.0
5.27

20.8
8.09
0.606

8,130

2,533

66.5oN

Summer solstice
Equinox
Winter solstice

4 months
01-Jun to 01-Oct
(122 d)

63
18.8
0.172

18.6
4.02
0.0026

5,649

1,589

Fig. 2. Plot of the waiting-in-line function (Eqs. 4a,b) for an

optimum irradiance of 700 mol m2 s1 PPFD. The graph also
shows a plot of total photosynthesis by a photosynthetic surface
thick enough that it is able to utilize all available PPFD that is
useable for photosynthesis (Eq. 7). Both equations have been
scaled for Pmax = 100%. Pg gross photosynthesis; PPFD
photosynthetic photon flux density.

A table was compiled of global horizontal irradiance

PPFD (mol m2 s1) on the ground for solar elevation
angles from 0 to 90 degrees plus 23.5 and 66.5 degrees.

602

A plot was made of PPFD vs. solar elevation angle. This

curve was found to fit a 4th order polynomial very well
(I = 0.00005643 4 0.0132731 3 + 0.805050 2 +
18.417200 + 20.217710, r = 0.99979, where is the
solar elevation angle in degrees) and so total PPFD
irradiance on flat horizontal ground can be calculated
easily for any solar elevation angle by a simple formula.
This could be used to calculate PPFD irradiance for any
given latitude, time and date of a year given the solar
elevation angle data from NREL Solar Radiation
Laboratory (2008).
Fig. 3A shows diurnal light curves for irradiance on a
flat planar surface for 37oN at summer solstice, winter
solstice, the two equinoxes and for an arbitrary date
14-Aug-2009. The total daily PPFD irradiance (mol m2)
can be calculated from the sum of the irradiance over the
course of a day using the trapezium rule and numerical
integration. The polynomial described above was used to
calculate PPFD at 15-min intervals during the course of
the day. The diurnal curve for 14-Aug-2009 lies between
the light curve for the summer solstice and the equinox. If
a similar plot is drawn for 37oS the photoperiods for the
solstices and equinoxes are almost the same as for 37oN
(there is a small effect due the small eccentricity of the
earths orbit). Total daily irradiance at the summer
solstice (21-Jun-2009) was very high (66.3 mol m2 d1)

PPFD AND PHOTOSYNTHESIS

Fig. 3. A: Diurnal light curves for irradiance on a flat planar

surface for 37o N at summer solstice, winter solstice, the two
equinoxes and for an arbitrary date 14-Aug-2009. Irradiances
are corrected for the thickness of the atmosphere through which
the sunlight passes. The maximum noon PPFD is 2,130 mol
m2 s1 (66.3 mol m2 d1) at the summer solstice (21-Jun-2009)
and falls to 970 mol m2 s1 (20.2 mol m2 d1) for the winter
solstice (21-Dec-2009). All mid-latitude diurnal light curves are
similar in shape to Fig. 3A. B: The estimated diurnal gross
photosynthesis (Pg) calculated using Eq. 7. In reality, the Pg
value for the winter solstice would be negligible because it
would be outside the growing season.

because of the very long day length at that latitude at the

summer solstice even though the maximum irradiance on
the 21-Jun-2009 solstice (PPFD = 2,130 mol m2 s-1)
was less than the maximum found at the Equator
(Table 1). Fig. 3B shows the estimated Pg in g(C) m2 s1
(Eq. 7) over the course of a day. The daily photosynthesis
curves are not the same shape as the irradiance curves
because of the saturating response of photosynthesis to
irradiance (Fig. 2).
Fig. 4A shows the daily irradiances for 37oN and S
calculated for each day of 2009. The curves are
noticeably flattened in the winter months because of
shorter daylight hours and lower solar elevation angles.
Total annual irradiance can be calculated by summation
but for a mid-latitude situation it is not realistic to sum Pg
over the entire year. For 37o latitude it would be reasonable to consider a growing season of about 7 months,
starting from the start of the month of the spring equinox
to the end of the month of the autumn equinox (Northern
hemisphere; 1 March to 1 October and southern hemisphere; 1 September to 1 April). The total growing-

Fig. 4. A: Total daily gross photosynthesis (Pg) on a flat planar

surface over a year for 37oN and S, calculated for each day of
2009. The curves are noticeably flattened in the winter months
because of short daylight hours and low solar elevation angles.
B: The estimated diurnal gross photosynthesis over a growing
season from 01 March to 01 October calculated using the sum
of Pg (estimated at 15-min intervals) over the course of each
day.

season irradiances for 37oN are 12,208 mol m2 (215

days) and 11,915 mol m2 (212 days) for 37oS. Plots of
daily Pg for days during the growing season are shown in
Fig. 4B. Similar annual irradiance curves can be
calculated for either hemisphere and for any latitude from
data on solar elevation angle (Gueymard 1995, Gueymard
2001, NREL Solar Radiation Laboratory 2008, SMARTS
2009).
For graphs of daily irradiances for the tropics of
Cancer and Capricorn and for the equator calculated daily
over the course of a solar year please see the
Supplementary material. Darwin (Northern Territory,
Australia: 1228S) was included as an example of an
intermediate site between the tropics of Capricorn and
Cancer but not on the equator. The curves are similar for
those found in mid-latitudes for the tropics of Cancer and
Capricorn but at the equatorial maximum irradiance
occurs twice a year at the equinoxes and is least when the
sun is directly over the tropics of Cancer and Capricorn.
The equator does not experience the maximum daily
irradiance found on earth; these are experienced at the
summer solstices at mid latitudes because of the longer
day-length. Darwin is located almost half way between
the equator and the tropic of Capricorn. The daily

603

R.J. RITCHIE

Fig. 5. Daily irradiances vs. gross photosynthesis (C3) for the

equator, tropics of Cancer or Capricorn (23o27 latitude) and
37o, 55o, and 66o27 latitudes. The curves for the equator and
latitudes up to 37o overlap and can be collectively described by
a single simple 3rd order polynomial of the form y =
0.00004574 x3 + 0.00591451 x2 + 0.148612 x where x is the
total daily PPFD in mol m2 d1, r = 0.9996.

irradiance at the equinoxes and the summer solstice are

almost equal and are not the dates at which maximum
irradiance occurs. Maximum irradiances (2,224 mol
m2 s1; 59.6 mol m2 d1) occur at midday on
16-Feb-2009 and 25-Oct-2009 when the sun past directly
overhead at the latitude of Darwin. There is little
variation in the day length over the year (about 11 to

13 h). Eq. 7 could be used to estimate Pg over the course

of a day and hence estimate daily carbon fixation.
In the case of tropical environments, it is realistic to
sum total irradiance over the year, because it is usually
warm enough for plants to grow all year round. Table 1
shows that the total annual irradiances in the tropics are
remarkably uniform: tropic of Cancer, 18,615 mol m2;
equator, 20,238 mol m2 and tropic of Capricorn, 18,425
mol m2, Darwin (12o28S), 19710 mol m2. Darwin
experiences a very high total annual irradiance even
though it does not experience the highest daily irradiance
on a global scale. Leaving aside water limitations,
potential daily Pg would be expected to closely follow
daily irradiance in tropical regions (see Supplementary
material).
The large data sets of irradiance data and estimates of
Pg used to prepare graphs of daily Pg vs. Id can be used to
plot estimated daily Pg vs. daily irradiance of C3 plants
(Fig. 5). The curves fit a simple 3rd order polynomial very
well. Curves for latitudes from the equator to 37o overlap
(37oN, tropic of Cancer, equator, Darwin, tropic of
Capricorn, 37oS). For tropical and mid-latitudes a general
relationship between daily PPFD irradiance and estimated
Pg based on the present study is;
Pg = 0.00004574 Id3 + 0.00591451 Id2 + 0.148612 Id (8)
where r = 0.9996.

Discussion
Use of the SMARTS software (Gueymard 1995,
Gueymard 2001, SMARTS 2009) more easily allows
comparisons between harvesting solar energy using solar
panels and proposals to grow plants for production of
biofuels. Much of the meteorological and climatology
literature quotes PAR irradiance in W m2: the approximate conversion factor derived from the present study
was 1 W m2 PAR 4.556 mol m2 s1 PPFD. This
value was calculated by the author based upon a mean
calculated from PPFD irradiance at solar elevation angles
from 30 to 90o through the US Standard Atmosphere.
This is within 1% of the conversion factor (4.6) given by
Gensler (1984). My value is slightly lower than the value
of 4.6 given by Gensler (1984) because there is a slight
red shift at low solar angles because blue light is more
heavily absorbed/scattered than red light when passing
through a thick layer of the earths atmosphere.
The SMARTS software (Gueymard 1995, Gueymard
2001, SMARTS 2009) has been shown in the present
study to useable for both northern and southern
hemispheres. Calculations made in the present study
agree very well with irradiance models developed by
Walsby (1997) for total irradiances for given latitude but
Walsbys models do not offer spectral information and
irradiances were expressed in W m2.
Using a combination of data calculated using the
SMARTS software and the solar elevation angle data from

604

the NREL solar radiation laboratory website it is possible

to calculate PPFD vs. solar time for any latitude and date
allowing for the attenuating effects of the varying
thicknesses of the atmosphere the incoming solar
radiation passes through during the course of a day
(relative atmospheric mass, RAM) (Fig. 3A). The model
will not only be useful for modelling primary production
from natural vegetation, crops and algal ponds for biofuel
production but will lead to more realistic estimates of
photosynthetic efficiency of plant communities in
climatic and environmental studies. Estimates of total
daily irradiance for any latitude and date, by numerical
integration, makes it possible to calculate the maximum
theoretical total irradiance for a whole year or for a
growing season (Fig. 4A).
The photosynthetic model developed in the present
study can be used to make estimates of theoretical
maximum Pg for a given date, latitude and hemisphere,
taking the saturation and optimum irradiance properties
of C3 photosynthesis into account (Eqs. 3, 4, and 6b;
Table 1). Estimates of gross photosynthesis shown in
Fig. 3B, 4B, and Table 1 are based upon a closed canopy
of C3 leaves able to use all useable light and having an
optimum irradiance requirement of about 700 mol
m2 s1 (PPFD). Figs. 3A,B and Table 1 show that high
rates of productivity are possible at mid-latitudes [up to
about 22 g(C) m2 d1]. However, winters at 37oN are

PPFD AND PHOTOSYNTHESIS

sufficiently cold to prevent significant photosynthesis

during part of the year. Here I have assumed a growing
season of about 7 months (01 March to 01 October).
Annual gross photosynthesis could be no more than about
4.1 kg(C) m2 y1 or 41 tonnes per hectare per year
(Figs. 4A,B; Table 1). Allowances for respiration outside
the growing season would reduce annual net photosynthesis considerably from estimates of net photosynthesis
made during the growing season.
Given optimal light and unlimited other resources it is
possible to achieve annual gross photosynthetic rates of
about 6.0 to 6.9 kg(C) m2 y1 or 60 to 69 t ha1 year1
in the tropics in C3 plants (Table 1, for examples of
tropical irradiance curves see Supplementary material,
Figs. 1S, 2S). Net production would be considerably
lower because complete usage of available light can only
be achieved by having a leaf area index much greater
than unity. A large leaf area index implies a large
biomass of leaves that perform minimal photosynthesis
because they are usually in heavy shade but would make
a large contribution to respiration, hence cutting down
overall net photosynthesis. Leaf area index is finely
regulated by plants because excess leaves are costly in
terms of carbon for the plant to synthesise and to
maintain.
Taking the primary production values for the tropics
(Table 1) it is possible to calculate that the overall
efficiency of gross photosynthesis reaches about 2.8% in
terms of conversion of moles of PPFD quanta into moles
of carbon (Eq. 8, Fig. 5). This agrees well with actual
measurements in the laboratory (Richmond 1999,
Falkowski and Raven 2007, Waltz 2009). Somewhat
counter-intuitively Eq. 8 (daily total Pg vs. PPFD for
tropical latitudes and 37oN, S) shows that the maximum
photosynthetic efficiency of a canopy of C3 leaves,
absorbing all useable light, is maximal under the
maximum daily irradiance found in the present study
(67.8 mol m2 d1; efficiency 2.8%) and falls to 2.1% at
20.7 mol m2 d1. My estimates of maximum efficiency
are less optimistic than Zhu et al. (2008) ( 4% for C3
plants to 6% for C4 plants) because mine were calculated
on a quantum, rather than an energy basis, and because
mine are based on total daily irradiance rather than
optimum irradiance.
In high summer high latitudes such as 55oN and the
Arctic Circle receive as much PPFD per day as the
tropics but only for a short period of the year (Table 1).
Some of the reasons why high productivity can be found
in the short summers of very high latitudes on land and
in oceanic systems are the combination of very long
hours of daylight, hence minimal night-time respiratory
losses during the growing season, combined with a lower
maximum irradiance due to the low solar angle which
lessens photoinhibition.
Calculations of irradiance calculated in the present
study are of course for cloudless skies. Thus, any site on
earth will receive considerably less irradiance than the

maximum. Few data sets of actual PAR or PPFD measurements under natural conditions (rather than crude
estimates derived from other measurements) are available
and modelling PAR or PPFD under cloud cover is not
straightforward (Rubio et al. 2005, Olofsson et al. 2007).
Cloud cover approximately behaves in the 400700 nm
windows like a neutral density filter (they appear white or
grey). Clouds also scatter light to create a diffuse light
source. There is also an apparent blue shift in total
irradiance because diffuse light from the blue celestial
dome (diffuse horizontal irradiance) becomes a more
significant contributor to total irradiance under cloudy
conditions. The absorbance and light scattering properties
of high cirrus, cumulus and nimbus clouds are different.
The effects of clouds upon irradiance also differ with
solar elevation angle; for example, the probability of a
ray of sunlight passing through a cloud is less for high
solar elevation angles than for low solar elevation angles.
The degree of cloudiness and the type of cloud cover
varies greatly with geographic location. Information on
hours of sunshine and cloudiness can be found in the
meteorological records and in the climatological literature
and from satellite monitoring projects (e.g. Rossow and
Duenas 2004, Reikard 2009) and on the worldwide web
(for example for Australia: Climate graphs and mapsaverage daily sunshine hours: [http://www.bom.gov.
au/climate/averages/climatology/sunshine_hours/sunhrs.
html], worldwide: International Satellite Cloud Climatology project (ISCCP): [http://www.gewex.org/ isccp.
html]). Meteorological measurements of irradiance are
generally expressed in W m2 (radiometric units measured
using a pyranometer) and cover the near-UV, visible and
near infrared wavelengths of light (full sunlight broadband or shortwave irradiance is about 1,100 W m2 for a
window of 3004,000 nm) and so are not restricted to the
PAR window (full sunlight global PAR irradiance is
about 485 W m2). Since the absoption/scattering
properties of clouds for infrared and PAR windows are so
different, conversion factors for solar radiation reported
as shortwave irradiance to PAR are only approximate and
actual measurements of the conversion factor vary
considerably (from 0.27 to 0.48) depending upon the
actual meteorological conditions and solar angle (Rubio
et al. 2005, Olofsson et al. 2007). Furthermore, meteorological data sets are often further processed in ways
which restrict their usefulness for photosynthetic studies.
For example, hours of sunshine is defined by the hours
where the broadband irradiance is above 120 W m2. This
is a relatively high value in photosynthetic terms, equivalent to about 220 mol m2 s1 (PPFD) which is about
10% of full sunlight. Leaves of many C3 plants growing
in full sunlight would be capable of photosynthesis
of more than half of the Pmax under such conditions
(Eqs. 4a,b).
Thus, zero hours of sunshine does not imply zero
photosynthesis: substantial photosynthesis can occur on
an overcast day registering zero hours of sunshine. This

605

R.J. RITCHIE

is quite the reverse of what a nave understanding of the

hours of sunshine statistic would lead one to believe. In
the tropics, in the wet season, it is not unusual for days to
be overcast in excess of 10 h per day. A diffuse-source
PPFD of 220 mol m2 s1 or about 5 to 8 mol m2 d1
would qualify as a cloudy day with zero sunshine but
would provide enough photons to fix about 1 g(C) m2 d1
(Eq. 8): the actual figure would probably be higher
because in the wet season some photoacclimation would
be expected to the lower light conditions, there would be
less photoinhibition, less photorespiration and a diffuse
light source favours high photosynthesis (Friend 2001,
Myeni et al. 2007, Posada et al. 2009). Primary production during tropical wet seasons and tropical high-altitude
cloud forests, more or less permanently under cloud
cover, are both very high and plants are photoacclimated
to lower average irradiance than full sunlight and the leaf
area index is adjusted accordingly (Myeni et al. 2007,
Posada et al. 2009).
Cloudiness significantly affects potential photosynthesis. Darwin has a dry-monsoon climate and very high
clear-sky irradiances. It would seem a good site for
biofuel production. Clear-sky irradiances at the latitude of
Darwin are very high over much of the year (Table 1) but
cloud cover would have large effects during the hot wetseason (October to March). Using the Australian
Meteorological Office data referred to above, during the
winter drought day lengths are about 11 h but hours of
sunshine are in excess of 9 h (zero cloud-cover) and so
daily maximum gross photosynthetic rates would remain
at about 11 g(C) m2 d1 (Table 1). During the wet season,
daylight hours are about 13 hours but hours of sunshine
average only 8 h per day and so maximum daily Gross
Photosynthesis would be the sum of about 8 hours
photosynthesis in full sunshine plus 5 hours under cloud
or a total of about 13 g(C) m2 d1. These maximum
estimates of Gross Photosynthesis are close to the
experimentally observed maximum primary productivity

found in rainforests and crops [about 10 g(C) m2 d1]

even though productivity by terrestrial vascular plants is
often water-limited, which causes stomatal closure, hence
limiting CO2 fixation (Taize and Zeiger 2002).
Very large amounts of time and money have been
spent on the concept of using algal ponds or more
conventional crops to produce biofuels (Richmond and
Zhou 1999, Antoni et al. 2007, Grobbelaar 2007, Huntley
and Redalje 2007, Waltz 2009). More detached assessments (Sheehan et al. 1998, Walker 2009, Larkum 2010)
have concluded that they are unlikely to be viable.
Walker (2009) points out that there seems to be a stubborn resistance to accepting the point that such schemes
have little chance of being viable. Some carbon fixation
rates claimed for algal production ponds [40 to 100 g(C)
m2 d1: Richmond and Zhou 1999, Waltz 2009] are well
above the theoretical limits calculated in the present study
for a photosynthetic organism (Eqs. 2, 3).
Some of the fundamental errors common in studies of
the relationship between irradiance and photosynthesis
and hence primary productivity are: (1) the simple fact
that sunlight is a dilute energy source is not understood,
(2) the quantum nature of the light reactions of
photosynthesis is not appreciated, (3) only part of the socalled PPFD spectrum is actually used for photosynthesis,
(4) the wrong solar irradiance units of measurement are
used or (5) global horizontal irradiance PPFD solar
irradiances for noon at the equator at equinox (2,220
mol m2 s1) are used where it is not appropriate to do so
(solar elevation angle is neglected), (6) the saturable
nature of the photosynthetic apparatus is not appreciated
(Ritchie 2008) and (7) photosynthetic efficiency under
natural lighting conditions is grossly overestimated
because photosynthetic efficiencies calculated from the
initial slope of P vs. I curves have been inappropriately
used (Ritchie 2008). The Excel routines used in the
present study are available from the author upon request.

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R.J. RITCHIE

Appendix: Configuration settings used for running SMARTS 2.9.5 software

Card 1: Comments
Enter a title name for SMARTS configuration.

Card 10: Regional albedo (predominate within r = 10 km)

Selected vegetation: Grazing field (unfertilised)

Card 2: Site pressure

Used standard default settings.
1013.25 Site pressure [mb]
0 Altitude at ground [km]
0 Height above ground [km]

Card 10a: Tilt albedo

Tilted surface & local albedo (predominate within r = 100 m)
Standard Default is for a tilted panel 37o from horizontal.
Selected tilt [deg] = 0 and azimuth [deg] = 0
Selected vegetation: Grazing field (unfertilised)

Card 3: Atmosphere
* For PAR spectra used appropriate latitudespecific atmosphere.
* For daily and annual irradiance selected: U.S.
Standard Atmosphere
* The effect of the difference between using the
default U.S. standard atmosphere and latitudespecific atmosphere is < 0.2% on PAR total.

Card 11: Spectral range

Selected spectral range:
minimum 400 nm: maximum 700 nm
Standard default solar constant [W m2]: 1,336.1
Standard default solar distance correction factor: 1.0

Card 4: Water vapour

Used standard default setting: Calculate from
reference atmosphere and altitude
* This option automatically uses the settings
specific for the atmosphere chosen on Card #3.

Card 12: Output

Output File options: create .OUT and .EXT files include spectral
results in both files
Spectral range to be printed [nm]: minimum 400 nm: maximum 700
nm: interval (step) 1
Spectral results:
Results in W m2 (PAR)
Direct normal irradiance
Diffuse horizontal irradiance
Global horizontal irradiance
Results in umol m2 s1 (PPFD)
Global horizontal photosynthetic photon flux
Direct normal photosynthetic photon flux
Diffuse horizontal photosynthetic photon flux

Card 5: Columnar ozone abundance

Used standard default setting: Use default from
reference atmosphere

Card 13: Circumsolar

Circumsolar calculations
Used standard setting: Bypass

Card 6: Gaseous absorption and pollution

used standard default setting: use defaults from
selected atmosphere

Card 14: Smoothing

Extra scanning/smoothing
Used standard setting: Bypass

Card 7: Carbon dioxide

Warning: Default value is out-of-date.
Used an estimated 2009 CE value of 389 (ppmv)
(NOAA Earth System Research Laboratory).

Card 15: Illuminance

Extra illuminance and photosynthetically active radiation
calculations
Used standard setting: Bypass

Card 7a: Extraterrestrial spectrum

Used standard default setting:
Gueymard 2004

Card 16: UV
Extra UV calculations
Used standard setting: Bypass

Card 8: Aerosol model

Used standard default setting:
Shettle & Fenn category: Rural

Card 17: Solar geometry

Solar position and air mass
* Two configurations used (some settings override Cards 1-16)
Configuration #1
Selected: Input elevation and azimuth angles [deg]
Two runs needed. Record numbers 1-60 and 61-90.
Apparent elevation angles (deg): 0-90 and azimuth angle (deg): 180
Configuration #2
Selected: Input year, month, day, hour, latitude, longitude, and
time zone
Specified year, month, day, hour and latitude of interest.
For the purposes of the present study the longitude was set at long.:
0 and time zone: 0

Card 9: Atmospheric turbidity

Used standard default turbidity: 0.084
Specified as: Default value:
Aerosol optical depth at 500 nm

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PPFD AND PHOTOSYNTHESIS

Supplementary figures

Fig. 1S. A: The diurnal irradiance patterns are quite different in

tropical locations because the sun passes directly overhead
twice a year. Light curves for Darwin, NT, Australia (latitude
12o28 S) for the summer and winter solstices, and the
equinoxes and at an arbitrary date of 14-Aug-2009. There is
little variation in the day-length over the year. The maximum
noon PPFD is 2,152 mol m2 s1 (59.9 mol m2 d1) at the
summer (wet-season) solstice (21-Dec-2009) and falls to 1,759
mol m2 s1 (43.4 mol m2 d1) for the winter (dry-season)
solstice (21-Jun-2009). B: the estimated gross photosynthesis
(Pg) (Eq. 7).

Fig. 2S. A: Daily irradiances for the tropics of Cancer and

Capricorn, the equator and for Darwin, NT, Australia (latitude
12o28 S). The curves are similar for those found in midlatitudes for the tropics of Cancer and Capricorn but at the
equator maximum irradiance occurs twice a year at the
equinoxes and is least when the sun is directly over the tropics
of Cancer and Capricorn. B: The estimated diurnal gross
photosynthesis (Pg) (Eq. 7).

609