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doi: 10.1111/j.1461-0248.2004.00567.x
REPORT
Goran I. Agren
Department of Ecology and
Environmental Research,
Swedish University of
Agricultural Sciences, PO Box
7072, SE-750 07 Uppsala,
Sweden
E-mail: goran.agren@eom.slu.se
Abstract
Evolution has set biochemical constraints on the chemical composition of living
organisms. These constraints seem to lead to increases in N : C and P : C ratios with
increasing relative growth rate for all types of organisms. The N : P ratio also seems to
decrease with relative growth rate for heterotrophs whereas autotrophs may show a
more complex behaviour. Here I will show that, from biochemical considerations, N : C
should increase linearly and P : C quadratically with relative growth rate in autotrophs
with the consequence that N : P increases at low relative growth rates, passes a
maximum and then decreases at high relative growth rates. These predictions are verified
against observations for a freshwater alga (Selenastrum minutum) and a tree seedling (Betula
pendula). Changes in temperature, light or other factors that affect the growth rate of
autotrophs interact with nutrient supply in such a way that there are no simple rules for
as to how N : P will change.
Keywords
Betula pendula, nitrogen, phosphorus, relative growth rate, RNA, Selenastrum minutum,
specific growth rate, stoichiometry.
Ecology Letters (2004) 7: 185191
INTRODUCTION
186 G. I. Agren
N N p bN C
dC
/CN Np
dt
dNp
/NP Pri
dt
l
bN
/CN
rPC l
1
l2 bP
/CN /NP
THEORY
Selenastrum minutum
Betula pendula
400
100
N limited
75
200
50
100
25
0
100
75
10
50
5
25
Redfield ratio
0
40
15
N : P (g g1)
30
10
20
10
0
0.0
N : P (g g1)
P : C (mg g1)
P : C (mg g1)
P limited
N : C (mg g1)
N : C (mg g1)
300
0.5
1.0
(day1)
1.5
0.00
0.05
0.10
0.15
0.20
0.25
0
0.30
(day1)
188 G. I. Agren
S. minutum
B. pendula
bN (mg g)1)
bP (mg g)1)
lmax
0.0101
0.00318
8.55
2.25
48.4
8.441
3.69
1.84
0.258
0.091
NP
(day)1)
rNP(lmax
NP)
(mg mg)1)
16.6
12.4
Table 2 Comparison of adjusted r2 for linear and quadratic regressions of N : C and P : C on relative growth rate. The number of
observations is given after the r2-value. The largest r2-values for N and P limited growth, respectively, are marked in bold
N limited
P limited
Species
Linear
Quadratic
Linear
Quadratic
Source
Selenastrum minutum
Betula pendula
0.939/18
0.931/10
0.897/18
0.896/10
0.826/21
0.831/16
0.898/21
0.881/16
Eucalyptus globulus
Scenedesmus acutus
Cyclotella meneghiana
Cyclotella meneghiana*
Aureoumbra lagunensis
0.981/9
0.955/6
0.924/6
0.853/6
0.902/5
0.958/9
0.888/6
0.974/6
0.779/6
0.983/5
0.830/8
0.831/6
0.826/6
0.897/4
0.957/4
0.890/8
0.704/6
0.938/6
0.943/4
0.900/4
*The values from the three lowest relative growth rates with P limitation were excluded from the analysis because of decreasing P : C with
increasing relative growth rate.
bNri
1
l2
l bN
/CN
/CN /NP
10
The equation for rPC remains unchanged. The consequences of introducing this additional N pool is thus to also
make the rNC function quadratic in l. However, the term
bNril//NP is at most c. 0.2 and the quadratic term will
probably not be distinguishable from the linear term with
current quality of data.
The numerical values for the parameters found with data
analysed in this paper all point to case 3, i.e. there should be
a clear maximum in the relationship between N : P and
relative growth rate. This is in contrast to what seems to be
the case for heterotrophs, where N : P decreases monotonically with relative growth rate. The parameters describing the interactions between the elements (/CN and /NP)
can also be compared with what can be predicted from
biochemical properties. Agren (1985b) estimated from
observed rates of the catalysing capacity of Rubisco that
/CN should have a value of 0.02 gC (mgN))1 day)1, which
is just a factor of 2 higher than observed for S. minutum.
Sterner & Elser (2002) give values for the rate of protein
synthesis by ribosomes. From their values we get /NP of
1.1 mgN (mgP))1 day)1. This is a factor of two lower than
the value given in Table 1 for B. pendula and a factor of
seven lower for S. minutum. Given the crude approximations
used to arrive at these estimates, the agreement between the
values obtained from biochemical considerations and whole
organism observations is not unreasonable, although this
point deserves further evaluation.
20
20
CN /2
15
Betula pendula
15
10
10
5
0
0.0
CN /2 & NP /2
0.5
1.0
1.5
0.0
0.1
0.2
N : P (g g 1)
N : P (g g 1)
Selenastrum minutum
5
0
0.3
190 G. I. Agren
Shafik, H.M., Herodek, S., Presing, M., Vorors, L. & Balogh, K.V.
(1997). Growth of Cyclotella meneghiniana Kutz. II. Growth and
cell composition under different growth rates with different
limiting nutrient. Ann. Limnol., 33, 223233.
Sterner, R.W. (1995). Elemental stoichiometry of species in ecosystems. In: Linking Species and Ecosystems (eds Jones, C.G. &
Lawton, J.H.). Chapman & Hall, New York, pp. 240252.
Sterner, R.W. & Elser, J.J. (2002). Ecological Stoichiometry The Biology
of Elements from Molecules to the Biosphere. Princeton University
Press, Princeton.
Terry, K.L. (1980). Nitrogen and phosphorus requirements of
Pavlova lutheri in continuous culture. Bot. Mar., 23, 757764.
APPENDIX. DERIVATION OF EQNS 5 AND 6
/NP Pri bN
bN
dt
dt
dt
dt
dC
/NP P bP C bN
dt
Terry, K.L., Laws, E.A. & Burns, D.J. (1985). Growth rate variation in the N:P requirement ratio of phytoplankton. J. Phycol., 21,
323329.
Editor, J. Elser
Manuscript received 22 October 2003
First decision made 2 December 2003
Second decision made 17 December 2003
Third decision made 23 December 2003
Manuscript accepted 5 January 2004
A2