Tree Physiology 24, 447–460
© 2004 Heron Publishing—Victoria, Canada
Terrestrial plants require nutrients in similar proportions
MAGNUS F. KNECHT1,2 and ANDERS GÖRANSSON3
1
Department of Ecology and Environmental Research, P.O. Box 7072, SLU, 750 07 Uppsala, Sweden
2
Corresponding author ([email protected])
3
Division of Education, P.O. Box 7070, SLU, 750 07 Uppsala, Sweden
Received June 13, 2003; accepted August 29, 2003; published online February 2, 2004
Summary Theoretical considerations based on nutrition experiments suggest that nutrient ratios of terrestrial plants are
similar to the Redfield ratio found in marine phytoplankton.
Laboratory experiments have shown that seedlings of many
different plant species have similar nutrient concentration ratios when supplied with nutrients at free access. However, at
free access, nutrients are likely to be taken up in amounts in excess of a plant’s requirements for growth. In further experiments, therefore, the supply rate of each nutrient was reduced
so that excessive uptake did not occur. Again, similar nutrient
ratios were found among the plant species tested, although the
ratios differed from those found in plants given free access to
nutrients. Based on the law of the minimum, we suggest that
optimum nutrient ratios be defined as the ratios found in plants
when all nutrients are limiting growth simultaneously. The literature on nutrient concentrations was surveyed to investigate
nutrient ratios in terrestrial ecosystems. Nutrients taken into
consideration were nitrogen, phosphorus, potassium, calcium
and magnesium. Based on the assumption that nitrogen is either the limiting nutrient or, when not limiting, is taken up only
in small excess amounts, we calculated nutrient ratios from
published data. The calculated ratios corresponded closely to
the ratios determined in laboratory and field experiments.
Keywords: ecosystems, free access nutrition, law of the minimum, nutrient ratios, plants.
Introduction
The element composition of organic matter has been a subject
of interest for centuries. Plants require some elements in large
amounts and others in only small amounts. Although there
may be significant differences among species, we predict that
individuals of the same species have more or less identical requirements. We also predict similarities among plant species
that have developed mechanisms for survival in similar environments.
Dry mass of terrestrial plants consists of about 450 mg g –1
carbon, 400 mg g –1 oxygen and 50 mg g –1 hydrogen, and the
concentrations of other elements vary within the remaining
50 to 100 mg g –1. Despite variability in concentrations of these
elements among species, patterns can be detected. For exam-
ple, seedlings of many species have an N:P mass ratio of
around 10:1 (whole plants), regardless of whether nutrients or
light limit plant growth (e.g., Ågren 1988, Ericsson 1994a,
Lambers et al. 1998). Garten (1976) found the same ratio for
plants sampled in the field. In many samples of marine
phytoplankton taken from various localities, Redfield (1958)
found that atoms of carbon, nitrogen and phosphorus were
generally present in the ratio 106:16:1 (i.e., 568:100:14) on a
mass basis. (Hereafter we present all ratios on a mass basis (as
a % of nitrogen) with nitrogen, set to 100, as the reference.)
However, these ratios do not necessarily correspond to a perfect nutrient balance. Geider and La Roche (2002) argue that a
realistic range for the critical N:P ratio may lie between 15 and
30, which, on a mass basis, is 100:7.4–14.8.
In terrestrial plants, concentrations of carbon, oxygen and
hydrogen are about the same (e.g., Salisbury and Ross 1992,
Lambers et al. 1998) and are therefore omitted from our investigation. If carbon, oxygen and hydrogen are excluded from
analysis, can we expect to find standard ratios between the
other essential elements in terrestrial plants? Because basic
physiological functions like photosynthesis and respiration involve similar molecules, enzymes and proteins, we predict
general patterns of occurrence of these elements, particularly
between the elements that are often referred to as nutrients.
Sterner and Elser (2002) have extensively investigated biochemical and cellular aspects of the N:P ratio and have provided mechanistic support for the existence of such patterns. A
complication in the analysis of nutrient ratios is that plants
may take up nutrients in excess of their requirements for
growth. For this reason, a basic statistical analysis of nutrient
ratios would be difficult to interpret. Instead, we will first
evaluate theoretically what we should expect.
Plant growth depends on the accessibility of essential nutrient elements. For a plant to achieve a certain relative growth
rate, RG (day –1), the minimum amount of all essential nutrient
elements required to maintain this rate must be available for
uptake. Because availability of all elements corresponding exactly to the plant’s requirements is unlikely, RG is instead limited by the availability of the nutrient element least available
relative to the plant’s requirement. This is generally referred to
as Liebig’s law of the minimum (Liebig 1840, 1855), although
448
KNECHT AND GÖRANSSON
it had already been formulated by Sprengel in 1828 (van der
Ploeg et al. 1999). The concentration of the limiting nutrient,
cn, in the plant can be considered a measure of the availability
of that nutrient.
Laboratory experiments have confirmed that certain minimum concentrations of some nutrients are required in the plant
before growth can occur (Ingestad 1982, 1987, Ingestad and
Lund 1986, Göransson 1993, 1994, Ericsson 1994a, Ågren
and Bosatta 1998). This amount has been defined as the minimum concentration of the nutrient, cn,min, and these amounts
are referred to collectively as “structural amounts,” indicating
that small amounts are required in structural compounds or for
trigger mechanisms (Figure 1) (Göransson 1993, Ågren and
Bosatta 1998). In this paper, the concentration range below
cn,min denotes the deficiency range. It has also been shown that
plant RG reaches its maximum at a specific nutrient concentration cn,opt (Ingestad 1979, Ingestad and Lund 1979, Ericsson
and Ingestad 1988, Ericsson and Kähr 1993, Ågren and
Bosatta 1998, Göransson 1999). A further increase in the nutrient concentration above cn,opt will not increase RG. We define
the concentration range between cn,min and cn,opt as the response
range and concentrations above cn,opt as the sufficiency range.
It has also been shown for several of the essential nutrients that
RG increases linearly with nutrient concentration within the response range (Ingestad 1979, Ingestad and Lund 1979,
Ericsson and Ingestad 1988, Ericsson and Kähr 1993, Ågren
and Bosatta 1998). We denote the slope of the response curve
by n-productivity, Pn, where n is the nutrient in question. Finally, at high concentrations, cn,tox, nutrients become toxic, the
growth rate decreases and the plants may eventually die. We
refer to concentrations above cn,tox as the toxicity range. The
mathematical expression for plant growth is then:
[0, c n,min ]

0


R G = (c n )Pn (c n − c n,min ) [c n,min, c n,opt ]

P (c
 n n,opt − c n,min ) [c n,opt , c n,tox ] 
(1)
A scenario where more than one nutrient is limiting growth
at the same time is still in agreement with the law of the minimum and can be demonstrated as follows. Assume that a plant
is limited to an RG by the internal concentration of nutrient A
(cA ) and that cA remains constant. Now, for another nutrient B,
let the internal concentration, cB, of the plant decrease (this can
happen if the availability of nutrient B decreases). Then at
some point, nutrient B will become limiting instead of A. At
the point where the limitation switches from nutrient A to B,
plant growth is limited by both nutrients simultaneously. This
condition can be extended to all essential nutrients, and we define optimum nutrient ratios as the ratio of nutrients when they
all limit growth simultaneously. At optimum nutrient ratios,
the internal concentration of each nutrient in the plant corresponds exactly to the current RG. The aim of our study was to
develop further insight into the relationships between nutrients
in terms of nutrient ratios.
We hypothesized that optimum nutrient ratios in terrestrial
plants are similar for a wide range of species. We used published data to test our hypothesis in an analysis based on optimum ratios that we determined in laboratory experiments and
that were later verified in a field experiment on mature trees. A
further assumption for our analysis was that nitrogen is generally taken up in amounts corresponding to the current RG of the
plant, or in amounts only slightly in excess of this value.
Methods
Figure 1. Definitions of growth responses in relation to nutrient concentrations in plants. The minimum concentration (cn,min ) of the nutrient is denoted “structural amount,” indicating that small amounts are
required in structural compounds or to trigger mechanisms. The concentration range between cn,min and cn,opt is the response range, and
concentrations above cn,opt are in the sufficiency range. At high concentrations (cn,tox ), nutrients become toxic and the growth rate decreases. We define this as the toxicity range.
The literature was searched for information about nutrient
concentrations of terrestrial plants growing under field conditions (see Appendix). The aim was to select nutrient data from
plants that had acclimated to the nutrient conditions at the site
where they were growing, thereby being close to a steady state.
Plants that had been fertilized were excluded. The data we required had to represent measurements of concentrations of
several nutrients, among them nitrogen and phosphorus, and
preferably at least one more nutrient. The amount of published
data that could be used for our analysis was much smaller than
expected. Published data that were given to only one significant digit and data representing combined values for all measured species in a stand were excluded. Data were available
either as mass per dry mass or mass per land area. In most
cases, the nutrient amounts in different plant parts were measured. We have, therefore, divided the data set into foliage
(leaves and needles), branches, stems, bark, roots and a group
referred to as other parts. When the plant parts from which the
samples were taken were not specified, data for the samples
were added to the foliage group, mainly because foliage was
the plant part most commonly measured, but also because
these data sets resembled the foliage group more than any other. Branches consist of all woody parts that are not stems, making this group rather diverse. The root group, which was not
TREE PHYSIOLOGY VOLUME 24, 2004
NUTRIENT RATIOS IN TERRESTRIAL PLANTS
split into size classes, is similarly diverse. Data are mostly
from the northern hemisphere, but some data are from Australia and New Zealand. The elements most frequently included
in the measurements were nitrogen, phosphorus, potassium,
calcium and magnesium, whereas sulfur, iron and manganese
were measured in only a few cases, so only the first five elements were included in our analysis. The data originated from
several different investigations (a number not easily determined because a reference sometimes represented several investigations and sometimes the same measurements appeared
in different references). Sampling was not standardized so that
rigorous statistical analyses could not be applied. The variety
of species represented was large. The data were coarsely categorized into three plant types: conifers, deciduous trees and
non-woody species (referred to as herbaceous species). Sometimes data were given for a family and not for a particular species. Because families can consist of both woody and
non-woody species, these data could not easily be fit into any
of our categories, and were therefore included in the group we
considered most appropriate. The three groups were expected
to exhibit different utilization and nutrient storage abilities.
In several laboratory experiments where nutrients were supplied at free access, it was found that, regardless of plant
species, the ratios of macronutrient elements were similar
(Ericsson 1981, Ingestad and Stoy 1982, Ingestad and Kähr
1985, Ingestad and Ågren 1988). These ratios are presented in
Table 1. Linder (1995) used these ratios as target values for foliage concentrations in a long-term nutrition experiment in
Flakaliden in northern Sweden. These target values were
changed as new research gave better estimates of optimum nutrient ratios (Ericsson and Ingestad 1988, Ericsson and Kähr
1993, 1995, Göransson 1993, 1994, 1999, 2001). In this field
experiment, the growth of Norway spruce was dramatically increased by the supply of nutrients. At optimum nutrient ratios
in foliage, no leaching of nutrients from the site has been detected.
Nutrient ratios determined at free access are associated with
excessive uptake of one or more nutrients. Detailed experiments on birch seedlings considering phosphorus (Ericsson
and Ingestad 1988), potassium (Ericsson and Kähr 1993),
calcium (T. Ericsson, Swedish University of Agricultural Sci-
Table 1. Nutrient ratios derived from laboratory experiments. A single
asterisk denotes plants supplied with nutrients at free access (Ericsson
1995) and double asterisks denote birch seedlings supplied with nutrients such that no excessive uptake of phosphorus (P) (Ericsson and
Ingestad, 1988), potassium (K) (Ericsson and Kähr 1993), calcium
(Ca) (Ericsson, unpublished data) or magnesium (Mg) (Ericsson and
Kähr 1995) occurred.
Plant type
Deciduous plants *
Coniferous plants *
Herbaceous plants *
Birch **
449
ences, Uppsala, unpublished data) and magnesium (Ericsson
and Kähr 1995) have determined the lowest ratio of these nutrients to nitrogen that could sustain maximum growth (Table 1). Although optimum ratios have been determined, these
ratios still need to be verified by laboratory experiments.
Göransson and Eldhuset (2001) found that when all nutrients
were supplied at the calculated optimum ratios, Norway
spruce seedlings grew at a lower rate than when provided with
free access to nutrients. A likely explanation for this lies in the
nitrogen source, which had an effect on the root morphology
and partitioning of biomass. Nitrogen was supplied solely
as nitrate or as a mixture of ammonium and nitrate, and
Göransson and Eldhuset (2001) found that plants supplied
solely with nitrate had a lower maximum RG. Furthermore, nitrate accumulated in the nutrient solution.
Results
The concentrations (or amounts per forest area) of phosphorus, potassium, calcium and magnesium were plotted against
nitrogen concentrations (or amounts per forest area) for each
of the three plant categories and for each category of plant
parts. In each plot, we also added a line representing the nutrient ratio we considered optimum (see also Table 1). We show
here only combined plots of foliage data for deciduous and coniferous species together with data from herbaceous species
(Figure 2). The same general patterns were observed in most
plots. Only bark and root data deviated from this pattern in
some cases. One set of root data had low concentrations of potassium. Later measurements of potassium at the same stand
gave values almost tenfold higher (not included in our data set,
H. Majdi, Swedish University of Agricultural Sciences,
Uppsala, personal communication). Most values fell on the
optimum line for phosphorus, yielding an average N:P ratio of
about 100:10. Values were mostly above the optimum line for
potassium, and values below the line were still very close to it.
For calcium, only one value in the entire data set was below the
optimum line. For magnesium, only a few values were below
the optimum line, although they were still close to the line.
The distributions of both concentrations and ratios varied
among plant parts and also among elements. Figure 3 shows
the distribution of the entire data set as well as a line representing the nutrient ratios we considered optimum. In leaves, the
distributions were narrowest for P, closely followed by K,
whereas Mg and particularly Ca showed considerable variability. Variability among plant parts was also large. Concentrations were highest in foliage and in the group comprising other
parts, fruits, flowers and cones.
Discussion
Suggested optimum nutrient ratios
N
P
K
Ca
Mg
100
100
100
100
14.6
15.0
14.3
8
64.6
47.5
68.3
30
7.0
8.0
8.3
2
9.4
7.5
8.7
4
Our analysis of nutrient ratios in terrestrial plants shows that
the hypothesis that plants in general require nutrients in the
same ratios cannot be rejected. On the contrary, despite the variety of ratios that occur in nature, patterns were observed that
fit the predictions we made based on theoretical considerations
and laboratory results. Thus, the concept of optimum nutrient
TREE PHYSIOLOGY ONLINE at http://heronpublishing.com
450
KNECHT AND GÖRANSSON
Figure 2. Phosphorus, potassium, calcium and magnesium concentrations
versus nitrogen in foliage of coniferous
(䊉), deciduous (䊊) and herbaceous
(䉲) species as found in the literature
(see Appendix). The dashed lines represent the optimum nutrient ratio.
ratios is meaningful and may be useful for determining nutrient limitations. However, to test our hypothesis more rigorously, we would require additional information involving fertilization experiments to identify the limiting nutrient, as well
as more sophisticated biochemically based determinations of
optimum nutrient ratios and information on relative growth
rates together with nutrient concentrations.
We now consider some important components of the theory.
The term optimum nutrient ratios has been used with slightly
different meanings. We defined optimum nutrient ratios as the
ratios found in plants when all nutrients are limiting growth simultaneously. Although based on the empirical relationship
between plant growth (see Equation 1) and nutrient concentration, this definition is logically stringent. Ingestad (1979) defined optimum nutrient ratios as the ratios found in plants
growing at maximum relative growth rate (RG,max) at free access to all nutrients. The diagnosis and recommendation integrated system (DRIS) (Beaufils 1973, Walworth and Sumner
1987, Bailey et al. 1997, Sinclair et al. 1997) represents a definition similar to that of Ingestad, whereas the deviation from
Figure 3. Distributions of plant tissue
nutrient ratios based on data found in
the literature (see Appendix). In each
plot, the line denotes optimum nutrient
ratio. For coniferous species, n ranged
from 206 to 231, for deciduous species
from 428 to 517, and for herbaceous
species from 211 to 244. Some outliers
were omitted to simplify the graphs.
One of the data sources reported extremely low potassium concentrations
in coniferous species (see text for further details).
TREE PHYSIOLOGY VOLUME 24, 2004
NUTRIENT RATIOS IN TERRESTRIAL PLANTS
optimum percentage (DOP) (Montañés 1993, Sinclair et al.
1997) method relates to concentrations of each nutrient.
According to the definition of optimum ratios that we have
adopted, should we expect these ratios to be constant within
the same species? The mechanistic investigation by Sterner
and Elser (2002) gives us reason to believe that these ratios are
not constant. Several experiments have shown that RG is linearly dependent on the concentration of the limiting nutrient in
the response range. In experiments with birch seedlings, cn,min
and cn,opt differed for nitrogen and phosphorus so that the N:P
ratio varied from less than 100:11 at low RG up to 100:8 at high
RG (see Ingestad 1979, Ingestad and Lund 1979, Ericsson and
Ingestad 1988). Experiments on Eucalyptus globulus Labill.
also showed variability in the N:P ratio, increasing from 100:3
at low RG to 100:10 at high RG (Ericsson 1994b). Hence, the
optimum nutrient ratio of nitrogen to phosphorus is not constant, but depends on the RG of the plant or the nutrient supply
rate.
When interpreting the ratios found in published data we
must consider the possibility of uptake of nutrients in excess of
that required to support the current growth rate. Are all elements that are available in excess also taken up in excessive
amounts? Laboratory studies have shown that all elements can
be taken up in excess of requirements for growth. However,
plants also have the ability to down-regulate uptake so that
toxic values are not reached (e.g., Lambers et al. 1998). The
graphs showing P, K, Ca and Mg versus N (Figure 3) suggest
that down-regulation may be pronounced for N, P and K, less
pronounced for Mg and almost nonexistent for Ca.
The availability of a nutrient to a plant must also play an important role. The reason we rarely find high concentrations of
nitrogen is probably because it is often the nutrient limiting
growth (Vitousek and Howarth 1991), and even if another nutrient is limiting growth, nitrogen is still often available in
limited amounts. The same reasoning can be applied for phosphorus. Limitation by other nutrients is less common, but does
occur.
Is it possible to determine which nutrient is limiting plant
growth in a given situation? Methods used in nutrition research are sometimes unreliable or difficult to perform
(Göransson 2001). The results of fertilization experiments led
Sinclair and Park (1993) to identify an inadequacy in the principle of the minimum. They concluded that substitution might
occur between nitrogen and potassium. However, substitution
of nutrient elements is biologically difficult to explain. Consider plants at a site where they are limited in growth by a nutrient, e.g., nitrogen, and that another nutrient, e.g., potassium,
is available in slightly greater amounts than required for the
current RG. The availability of nutrients should be considered
in terms of the rates at which they can be taken up and the resulting concentrations in the plant. The relative uptake rate
(rate of uptake of the nutrient per amount of nutrient in the
plant) of each nutrient then has to be at least equal to RG if the
concentration of the nutrient is not to decrease. If nitrogen is
applied as fertilizer, it may be taken up at a higher rate and the
plants will have a higher RG, but only if the other nutrients can
451
be taken up at a rate at least corresponding to the new RG. If the
relative uptake rate of, for example, potassium is lower than
the new relative rate at which nitrogen can be taken up, we
must also expect an increased yield response when fertilizing
with a combination of nitrogen and potassium. Furthermore,
there may be an acclimation to the new nutrient availability; as
Sinclair and Park (1993) pointed out, the root/mass ratio (ratio
between root biomass and total plant biomass) decreases as nitrogen concentration in the plant increases, and this may lead
to decreased uptake capability for other nutrients. Furthermore, nutrients added to the soil may become immobilized before plants can take them up, thus reducing the effect of
fertilization.
The ratios calculated from published data agreed well with
the results obtained from experiments in climate chambers. In
particular, the mean N:P ratio in nature seems to be close to optimum. Most values were closely distributed around an N:P
value equal to 100:10 (see Figures 2 and 3). The extreme values occurred mostly when the concentration of either N or P
was low, suggesting either strong nitrogen or strong phosphorus limitation. This indicates that excessive uptake of nitrogen
or phosphorus generally occurred in small amounts and that
there is a relatively strong down-regulation restricting excessive uptake of the two elements when they are not limiting
growth. The concentrations of potassium, calcium and magnesium were, in most cases, higher than the optimum ratios, confirming that plants are generally limited by either nitrogen or
phosphorus. A comparison of ratios to N of K, Mg and Ca
showed that excessive uptake occurred in the order Ca > Mg >
K. The plots of Ca versus N suggest that Ca is generally taken
up in amounts corresponding to availability rather than to plant
requirements. This supports the hypothesis that calcium uptake depends partly on water flow into the plant. On the other
hand, both K and Mg were correlated with N, although it
seems that excessive uptake may occur, provided the optimum
ratios presented are correct. These two elements are generally
not in short supply, compared with nitrogen and phosphorus.
Koerselman and Meuleman (1996) suggested that the N:P
ratio may indicate the nature of nutrient limitation. They suggested that an N:P ratio less than 14 (100:7) by mass in wetland species indicates N limitation and an N:P ratio over 16
(100:6) indicates P limitation, based on the response to fertilization. This would indicate lower amounts of P required in
this type of ecosystem. In a recent paper referring to the same
investigation, Güsewell et al. (2003) found that nitrogen limitation could not clearly be shown for N:P ratios lower than 14
(100:7). These findings may be explained by the generally
high atmospheric nitrogen deposition rate in The Netherlands
where this investigation was made, and that nutrients other
than N or P may be limiting growth. The critical N:P ratio that
Geider and La Roche (2002) suggest for marine phytoplankton (100:7–100:13) is consistent with our results.
We suggest that future studies include laboratory experiments where plants growing at different RG are limited by several nutrients simultaneously. Laboratory experiments should
also be extended over a longer time period and throughout the
TREE PHYSIOLOGY ONLINE at http://heronpublishing.com
452
KNECHT AND GÖRANSSON
annual cycle. Field data should include determinations of the
limiting nutrient (or nutrients) and estimates of RG. Current
field experiments in progress in Sweden may provide these
data. If general optimum nutrient ratios in plants can be consistently determined, they may provide a means of optimizing
fertilization and minimizing the accumulation and leaching of
nutrients.
References
Ågren, G.I. 1988. Ideal nutrient productivities and nutrient proportions in plant growth. Plant Cell Environ. 11:613–620.
Ågren, G.I. and E. Bosatta. 1998. Theoretical ecosystem ecology. Understanding element cycles. Cambridge University Press, New
York, 250 p.
Alban, D.H., D.A. Perala and B.E. Schlaegel. 1978. Biomass and nutrient distribution in aspen, pine, and spruce stands on the same soil
type in Minnesota. Can. J. For. Res. 8:290–299.
Bailey, J.S., J.A.M. Beattle and D.J. Kildpatrick. 1997. The diagnosis
and recommendation integrated system (DRIS) for diagnosing the
nutrient status of grassland swards: I. Model establishment. Plant
Soil 197:127–135.
Beaufils, E.R. 1973. Diagnosis and recommendation integrated system (DRIS). Natal (South Africa): University of Natal. Soil Science Bulletin No. 1, 132 p.
Cole, D.W. and M. Rapp. 1981. Elemental cycling in forest ecosystems. In Dynamic Properties of Forest Ecosystems. Ed. D.E.
Reichle. Cambridge University Press, London, pp 341–409.
Cornelissen, J.H.C., M.J.A. Werger, P. Castro-Diez, J.W.A. van
Rheenen and A.P. Rowland. 1997. Foliar nutrients in relation to
growth, allocation and leaf traits in seedlings of a wide range of
woody plant species and types. Oecologia 111:460–469.
Day, F.P. and C.D. Monk. 1977. Seasonal nutrient dynamics in the
vegetation on a southern Appalachian watershed. Am. J. Bot.
64:1126–1139.
Ericsson, T. 1981. Effects of varied nitrogen stress on growth and nutrition in three Salix clones. Physiol. Plant. 51:423–429.
Ericsson, T. 1994a. Nutrient dynamics and requirement of forest
crops. N.Z. J. For. Sci. 24:133–168.
Ericsson, T. 1994b. Nutrient requirement of Eucalyptus globulus
seedlings. In Eucalyptus for Biomass Production. Eds. J.S. Pereira
and H. Pereira. Commission of the European Communities, 322 p.
Ericsson, T. and T. Ingestad. 1988. Nutrition and growth of birch
seedlings at varied relative phosphorus addition rate. Physiol.
Plant. 72:227–235.
Ericsson, T. and M. Kähr. 1993. Growth and nutrition of birch seedlings in relation to potassium supply rate. Trees 7:78–85.
Ericsson, T. and M. Kähr. 1995. Growth and nutrition of birch seedlings at varied relative addition rates of magnesium. Tree Physiol.
15:85–93.
Fahey, T.J., J.J. Battles and G.F. Wilson. 1998. Responses of early
successional northern hardwood forest to changes in nutrient availability. Ecol. Monogr. 68:183–212.
Foulds, W. 1993. Nutrient concentrations of foliage and soil in
South-western Australia. New Phytol. 125:529–546.
Garten, Jr., C.T. 1976. Correlations between concentrations of elements in plants. Nature 261:686–688.
Geider, R.J. and J. La Roche. 2002. Redfield revisited: variability of
C:N:P in marine microalgae and its biochemical basis. Eur. J.
Phycol. 37:1–17.
Göransson, A. 1993. Growth and nutrition of small Betula pendula
plants at different relative addition rates of iron. Trees 8:31–38.
Göransson, A. 1994. Growth and nutrition of small Betula pendula
plants at different relative addition rates of manganese. Tree
Physiol. 14:375–381.
Göransson, A. 1999. Growth and nutrition of Betula pendula at different relative supply rates of zinc. Tree Physiol. 19:111–116.
Göransson, A. 2001. A technique for quantitative trace element and
micronutrient studies in plants. In Trace Elements in the Rhizosphere. Eds. G.R. Gobran, W.W. Wenzel and E. Lombi. CRC Press,
Boca Raton, NY, pp 697–700.
Göransson, A. and T.D. Eldhuset. 2001. Is the (Ca + K + Mg)/Al ratio
in the soil solution a predictive tool for estimating forest damage?
Water Air Soil Pollut. 1:57–74.
Green, D.C. and D.F. Grigal. 1980. Nutrient accumulation in jack
pine stands on deep and shallow soils over bedrock. For. Sci. 26:
325–333.
Güsewell, S., W. Koerselman and J.T.A. Verhoeven. 2003. Biomass
N:P ratios as indicators of nutrient limitation for plant populations
in wetlands. Ecol. Appl. 13:372–384.
Helmisaari, H.S. 1990. Temporal variation in nutrient concentrations
of Pinus sylvestris needles. Scan. J. For. Res. 5:177–194.
Ingestad, T. 1979. Nitrogen stress in birch seedlings. II. N, K, P, Ca
and Mg nutrition. Physiol. Plant. 45:149–157.
Ingestad, T. 1982. Relative addition rate and external concentration;
driving variables used in plant nutrition research. Plant Cell Environ. 1:443–453.
Ingestad, T. 1987. New concepts on soil fertility and plant nutrition as
illustrated by research on forest trees and stands. Geoderma 40:
237–252.
Ingestad, T. and G.I. Ågren. 1988. Nutrient uptake and allocation at
steady-state nutrition. Physiol. Plant. 72:450–459.
Ingestad, T. and M. Kähr. 1985. Nutrition and growth of coniferous
seedlings at varied relative nitrogen addition rate. Physiol. Plant.
65:109–116.
Ingestad, T. and A.-B. Lund. 1979. Nitrogen stress in birch seedlings.
I. Growth technique and growth. Physiol. Plant. 45:137–148.
Ingestad, T. and A.-B. Lund. 1986. Theory and techniques for steady
state mineral nutrition and growth of plants. Scand. J. For. Res. 1:
439–453.
Ingestad, T. and V. Stoy. 1982. Mineral nutrition of wheat, rye, barley,
and oat seedlings in nutrient solutions. Swed. J. Agric. Res. 12:
185–192.
Johnson, F.L. and P.G. Risser. 1974. Biomass, annual net primary production, and dynamics of six mineral elements in a post oak–blackjack oak forest. Ecology 55:1246–1258.
Koerselman, W. and A.F.M. Meuleman. 1996. The vegetation N:P
ratio: a new tool to detect the nature of nutrient limitation. J. Appl.
Ecol. 33:1441–1450.
Lambers, H., F.S. Chapin, III and T.L. Pons. 1998. Plant physiological
ecology. Springer-Verlag, New York, 540 p.
Liebig, J. 1840. Die organische Chemie in ihrer Anwendung auf
Agricultur und Physiologie. Friedrich Vieweg und Sohn, Braunschweig, Germany, 352 p.
Liebig, J. 1855. Die Grundsätze der Agricultur-Chemie mit Rücksich
auf die in England angestellten Untersuchungen. 2nd Edn. Friedrich Vieweg und Sohn, Braunschweig, Germany, 107 p.
Linder, S. 1995. Foliar analysis for detecting and correcting nutrient
imbalances in Norway spruce. Ecol. Bull. 44:178–190.
Madgwick, H.A.I., P. Beets and S. Gallagher. 1981. Dry matter accumulation, nutrient and energy content of the aboveground portion
of 4-year-old stands of Eucalyptus nitens and E. fastigata. N.Z. J.
For. Sci. 11:53–59.
TREE PHYSIOLOGY VOLUME 24, 2004
NUTRIENT RATIOS IN TERRESTRIAL PLANTS
Majdi, H. and U. Rosengren-Brink. 1994. Effects of ammonium sulphate application on the rhizosphere, fine-root and needle chemistry
in a Picea abies (L.) Karst. stand. Plant Soil 162:71–80.
Miller, H.G., J.D. Miller and J.M. Cooper. 1980. Biomass and nutrient
accumulation at different growth rates in thinned plantations of
Corsican pine. Forestry 53:23–39.
Montañés, L., L. Heras, J. Abadía and M. Sanz. 1993. Plant analysis interpretation based on a new index: deviation from optimum percentage (DOP). J. Plant Nutr. 16:1289–1308.
Nihlgård, B. 1972. Plant biomass, primary production and distribution
of chemical elements in a beech and a planted spruce forest in south
Sweden. (Fagus sylvatica, Picea abies). Oikos 23:69–81.
Nihlgård, B. and L. Lindgren. 1977. Plant biomass, primary production
and bioelements of three mature beech (Fagus sylvatica) forests in
south Sweden. Oikos 28:95–104.
Odum, H.T. 1970. Summary: an emerging view of the ecological system at El Verde. In Tropical Rain Forest. Ed. R.F. Pigeon. U.S.
Atomic Energy Commision, Washington, DC, pp I191–I281.
Pope, P.E. 1979. The effect of genotype on biomass and nutrient content
in 11-year-old loblolly pine plantations. Can. J. For. Res. 9:224–230.
Redfield, A.C. 1958. The biological control of chemical factors in the
environment. Am. Sci. 46:205–221.
Rogers, R.W. and W.E. Westman. 1977. Seasonal nutrient dynamics of
litter in a subtropical eucalypt forest, North Stradbroke Island. Aust.
J. Bot. 25:47–58.
Rolfe, G.L., M.A. Akhtar and L.E. Arnold. 1978. Nutrient distribution
and flux in a mature oak–hickory forest. For. Sci. 24:122–130.
Salisbury, F.B and C.W. Ross. 1992. Plant physiology. 4th Edn.
Wadsworth Publishing, Belmont, CA, 682 p.
Schlesinger, W.H. 1978. Community structure, dynamics and nutrient
cycling in the Okefenokee cypress swamp-forest. Ecol. Monogr.
48:43–65.
Sinclair, T.R. and W.I. Park. 1993. Inadequacy of the Liebig limitingfactor paradigm for explaining varying crop yields. Agron. J. 85:
742–746.
Sinclair, A.G., J.D. Morrison, C. Smith and K.G. Dodds. 1997. Determination of optimum nutrient element ratios in plant tissue. J. Plant
Nutr. 20:1069–1083.
453
Sterner, R.W. and J.J. Elser. 2002. Ecological stoichiometry. The biology of elements from molecules to the biosphere. Princeton
University Press, Princeton, 439 p.
Tanner, E.V.J. 1977. Four mountain rain forests of Jamaica: a quantitative characterization of the floristics, the soils and the foliar
mineral levels, and a discussion of the interrelations. J. Ecol.
65:883–918.
Thompson, K., J.A. Parkinson, S.R. Band and R.E. Spencer. 1997.
A comparative study of leaf nutrient concentrations in a regional
herbaceous flora. New Phytol. 136:679–689.
Turner, J. 1981. Nutrient cycling in an age sequence of western
Washington Douglas-fir stands. Ann. Bot. 48:159–169.
Turner, J. and M.J. Singer. 1975. Nutrient distribution and cycling in
a sub-alpine coniferous forest ecosystem. J. Appl. Ecol. 13:
295–301.
Turner, J., D.W. Cole and S.P. Gessel. 1976. Mineral nutrient accumulation and cycling in a stand of red alder (Alnus rubra). J. Ecol.
64:965–974.
van der Ploeg, R.R., W. Böhm and M.B. Kirkham. 1999. History of
soil science: on the origin of the theory of mineral nutrition of
plants and the law of the minimum. Soil Sci. Soc. Am. J. 63:
1055–1062.
Vitousek, P.M. and R.W. Howarth. 1991. Nitrogen limitation on
land and in the sea: how can it occur? Biogeochemistry 13:
87–116.
Vitousek, P.M., G. Aplet, D. Turner and J.J. Lockwood. 1992. The
Mauna Loa environmental matrix: foliar and soil nutrients.
Oecologia 89:372–382.
Vitousek, P.M., D.R. Turner and K. Kitayama. 1995. Foliar nutrients
during long-term soil development in Hawaiian montane rain forest. Ecology 76:712–720.
Walworth, J.L. and M.E. Sumner. 1987. The diagnosis and recommendation integrated system (DRIS). Adv. Soil Sci. 6:149–188.
Whittaker, R.H., G.E. Likens, F.H. Bormann, J.S. Eaton and T.G.
Siccama. 1979. The Hubbard Brook ecosystem study: forest nutrient cycling and element behavior. Ecology 60:203–220.
Woodwell, G.M., R.H. Whittaker and R.A. Houghton. 1975. Nutrient concentrations in plants in the Brookhaven oak–pine forest.
Ecology 56:318–332.
TREE PHYSIOLOGY ONLINE at http://heronpublishing.com
454
KNECHT AND GÖRANSSON
Appendix
Table A1. Origin of nutrient data.
Reference
Species
Location
Alban et al. 1978
Alban et al. 1978
Alban et al. 1978
Alban et al. 1978
Cole and Rapp 1981
Cole and Rapp 1981
Cole and Rapp 1981
Cole and Rapp 1981
Cole and Rapp 1981
Cole and Rapp 1981
Cole and Rapp 1981
Cole and Rapp 1981
Cole and Rapp 1981
Cole and Rapp 1981
Cole and Rapp 1981
Cole and Rapp 1981
Cole and Rapp 1981
Cole and Rapp 1981
Cole and Rapp 1981
Cole and Rapp 1981
Cole and Rapp 1981
Cole and Rapp 1981
Cole and Rapp 1981
Cole and Rapp 1981
Cole and Rapp 1981
Cole and Rapp 1981
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Picea glauca
Pinus banksiana
Pinus resinosa
Populus tremuloides
Abies firma
Acer, Betula, Fagus
Alnus rubra
Fagus sylvatica
Fagus sylvatica
Liriodendron tulipifera
Picea abies
Picea abies
Picea abies
Picea mariana
Pinus echinata
Pinus strobus
Pseudotsuga menziesii
Pseudotsuga menziesii
Quercus ilex
Quercus prinus
Quercus–Betula
Quercus–Carya
Quercus–Carya
Quercus mixed
Tsuga heterophylla
Tsuga sieboldii
Acer campestre
Acer platanoides
Acer pseudoplatanus
Aesculus hippocastanum
Alnus glutinosa
Arbutus unedo
Atriplex halimus
Berberis vulgaris
Betula pendula
Buddleja davidii
Buxus sempervirens
Calluna vulgaris
Castanea sativa
Celtis australis
Cistus albidus
Cistus clusii
Cistus laurifolius
Cornus sanguinea
Crataegus monogyna
Cytisus scoparius
Daphne gnidium
Daphne mezereum
Dryas octopetala
Empetrum nigrum
Erica cinerea
Fagus sylvatica
Frangula alnus
Fraxinus excelsior
Hebe × franciscana
Hedera helix
Minnesota, USA
Minnesota, USA
Minnesota, USA
Minnesota, USA
Yusuhara, Japan
New Hampshire, USA
Washington, USA
Solling, Germany
Kongalund, Sweden
Tennessee, USA
Karelia, USSR
Kongalund, Sweden
Solling, Germany
Alaska, USA
Tennessee, USA
North Carolina, USA
Oregon, USA
Washington, USA
Rouquet, France
Tennessee, USA
Meathop Wood, U.K.
North Carolina, USA
Tennessee, USA
Virelles, Belgium
Oregon, USA
Yusuhara, Japan
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
Continued on facing page.
TREE PHYSIOLOGY VOLUME 24, 2004
NUTRIENT RATIOS IN TERRESTRIAL PLANTS
455
Table A1 Cont’d. Origin of nutrient data.
Reference
Species
Location
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Cornelissen et al. 1997
Day and Monk 1977
Day and Monk 1977
Day and Monk 1977
Day and Monk 1977
Day and Monk 1977
Day and Monk 1977
Helianthemum nummularium
Helianthemum squamatum
Hippophae rhamnoides
Ilex aquifolium
Juglans regia
Laburnum anagyroides
Larix decidua
Ligustrum vulgare
Linum suffruticosum
Lonicera implexa
Lonicera periclymenum
Malus sylvestris
Picea sitchensis
Pinus halepensis
Pinus sylvestris
Pistacia lentiscus
Prunus laurocerasus
Prunus lusitanica
Prunus spinosa
Quercus cerris
Quercus coccifera
Quercus faginea
Quercus ilex ballota
Quercus ilex ilex
Quercus petraea
Quercus pubescens
Quercus robur
Quercus rubra
Quercus suber
Rhamnus alaternus
Rhamnus cathartica
Rhamnus lycioides
Ribes nigrum
Ribes uva-crispa
Rosa arvensis
Rosmarinus officinalis
Rubus fruticosus
Rubus ulmifolius
Salix caprea
Sambucus nigra
Santolina chamaecyparissus
Solanum dulcamara
Sorbus aucuparia
Taxus baccata
Thymus polytrichus
Tilia cordata
Ulex europaeus
Ulex gallii
Ulmis glabra
Vaccinium myrtillus
Viburnum tinus
Viburnum opulus
Acer pensylvanicum
Acer rubrum
Amelanchier arborea
Betula lutea
Carya glabra
Castanea dentata
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
United Kingdom and northern Spain
North Carolina, USA
North Carolina, USA
North Carolina, USA
North Carolina, USA
North Carolina, USA
North Carolina, USA
Continued on next page.
TREE PHYSIOLOGY ONLINE at http://heronpublishing.com
456
KNECHT AND GÖRANSSON
Table A1 Cont’d. Origin of nutrient data.
References
Species
Location
Day and Monk 1977
Day and Monk 1977
Day and Monk 1977
Day and Monk 1977
Day and Monk 1977
Day and Monk 1977
Day and Monk 1977
Day and Monk 1977
Day and Monk 1977
Day and Monk 1977
Day and Monk 1977
Day and Monk 1977
Day and Monk 1977
Day and Monk 1977
Day and Monk 1977
Day and Monk 1977
Day and Monk 1977
Day and Monk 1977
Fahey et al. 1998
Fahey et al. 1998
Fahey et al. 1998
Fahey et al. 1998
Fahey et al. 1998
Green and Grigal 1980
Helmisaari 1990
Johnson and Risser 1974
Johnson and Risser 1974
Madgwick et al. 1981
Madgwick et al. 1981
Majdi and Rosengren-Brink 1994
Miller et al. 1980
Nihlgård 1972
Nihlgård 1972
Nihlgård and Lindgren 1977
Odum 1970
Pope 1979
Rogers and Westman 1977
Rogers and Westman 1977
Rolfe et al. 1978
Rolfe et al. 1978
Rolfe et al. 1978
Rolfe et al. 1978
Rolfe et al. 1978
Rolfe et al. 1978
Schlesinger 1978
Schlesinger 1978
Schlesinger 1978
Schlesinger 1978
Schlesinger 1978
Schlesinger 1978
Schlesinger 1978
Schlesinger 1978
Schlesinger 1978
Schlesinger 1978
Schlesinger 1978
Schlesinger 1978
Tanner 1977
Tanner 1977
Cornus florida
Hamamelis virginiana
Liriodendron tulipifera
Magnolia fraseri
Nyssa sylvatica
Other herbs
Oxydendrum arboreum
Pyrularia pubera
Quercus alba
Quercus coccinea
Quercus prinus
Quercus rubra
Quercus velutina
Rhododendron calendulaceum
Robinia pseudoacacia
Sassafras albidum
Symplocos tinctoria
Vaccinium stamineum
Acer pensylvanicum
Betula lutea
Betula papyrifera
Fagus grandifolia
Prunus pensylvanica
Pinus banksiana
Pinus sylvestris
Quercus marilandica
Quercus stellata
Eucalyptus fastigiata
Eucalyptus nitens
Picea abies
Pinus nigra
Fagus sylvatica
Picea abies
Fagus sylvatica
Not specified
Pinus taeda
Eucalyptus signata
Eucalyptus umbra
Acer saccharum
Carya glabra
Cornus florida
Quercus alba
Quercus rubra
Ulmus alata
Clethra alnifolia
Cyrilla racemiflora
Eriocaulon compressum
Ilex cassine
Itea virginica
Leucothoe racemosa
Lyonia lucida
Nyssa sylvatica var. biflora
Phoradendron serotinum
Taxodium distichum
Tillandsia usneoides
Usnea spp.
Alchornea latifolia
Blakea trinervia
North Carolina, USA
North Carolina, USA
North Carolina, USA
North Carolina, USA
North Carolina, USA
North Carolina, USA
North Carolina, USA
North Carolina, USA
North Carolina, USA
North Carolina, USA
North Carolina, USA
North Carolina, USA
North Carolina, USA
North Carolina, USA
North Carolina, USA
North Carolina, USA
North Carolina, USA
North Carolina, USA
White mountain National Forest, New Hampshire, USA
White mountain National Forest, New Hampshire, USA
White mountain National Forest, New Hampshire, USA
White mountain National Forest, New Hampshire, USA
White mountain National Forest, New Hampshire, USA
Minnesota, USA
Ilomantsi, Finland
Oklahoma, USA
Oklahoma, USA
Rotoehu State Forest, New Zealand
Rotoehu State Forest, New Zealand
South western Sweden
Culbin, Scotland (U.K.)
South Sweden
South Sweden
South Sweden
El Verde, Puerto Rico
Arkansas, USA
Queensland, Australia
Queensland, Australia
Illinois, USA
Illinois, USA
Illinois, USA
Illinois, USA
Illinois, USA
Illinois, USA
Georgia, USA
Georgia, USA
Georgia, USA
Georgia, USA
Georgia, USA
Georgia, USA
Georgia, USA
Georgia, USA
Georgia, USA
Georgia, USA
Georgia, USA
Georgia, USA
Blue Mountains, Jamaica
Blue Mountains, Jamaica
Continued on facing page.
TREE PHYSIOLOGY VOLUME 24, 2004
NUTRIENT RATIOS IN TERRESTRIAL PLANTS
457
Table A1 Cont’d. Origin of nutrient data.
References
Species
Location
Tanner 1977
Tanner 1977
Tanner 1977
Tanner 1977
Tanner 1977
Tanner 1977
Tanner 1977
Tanner 1977
Tanner 1977
Tanner 1977
Tanner 1977
Tanner 1977
Tanner 1977
Tanner 1977
Tanner 1977
Tanner 1977
Tanner 1977
Tanner 1977
Tanner 1977
Tanner 1977
Tanner 1977
Tanner 1977
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Chaetocarpus globosus
Clethra occidentalis
Cleyera theaeoides
Clusia cf. havetioides
Cyrilla racemiflora
Eugenia virgultosa
Eupatorium critoniforme
Guarea swartzii
Haemanthus incrassatus
Hedyosmum arborescens
Ilex macfadyenii
Ilex obcordata
Laplacea haematoxylon
Lyonia cf. octandra
Mecranium purpurascens
Meriania purpurea
Palicourea alpina
Podocarpus urbanii
Schefflera sciadophyllum
Solanum punctulatum
Turpina occidentalis
Vaccinium meridionale
Adoxa moschatellina
Agrostis capillaris
Alliaria petiolata
Allium ursinum
Anemone nemorosa
Angelica sylvestris
Anisantha sterilis
Anthoxanthum odoratum
Anthriscus sylvestris
Arabidopsis thaliana
Arrhenatherum elatius
Arum maculatum
Brachypodium pinnatum
Briza media
Bromopsis erecta
Calluna vulgaris
Campanula rotundifolia
Carex flacca
Catapodium rigidum
Catha palustris
Centaurea nigra
Centaurea scabiosa
Cerastium fontanum
Chamerion angustifolium
Chenopodium album
Chysosplenium oppositifolium
Conopodium majus
Conyza canadensis
Dactylis glomerata
Deschampsia caespitosa
Deschampsia flexuosa
Digitalis purpurea
Elytrigia repens
Epilobium hirsutum
Eriophorum angustifolium
Eriophorum vaginatum
Blue Mountains, Jamaica
Blue Mountains, Jamaica
Blue Mountains, Jamaica
Blue Mountains, Jamaica
Blue Mountains, Jamaica
Blue Mountains, Jamaica
Blue Mountains, Jamaica
Blue Mountains, Jamaica
Blue Mountains, Jamaica
Blue Mountains, Jamaica
Blue Mountains, Jamaica
Blue Mountains, Jamaica
Blue Mountains, Jamaica
Blue Mountains, Jamaica
Blue Mountains, Jamaica
Blue Mountains, Jamaica
Blue Mountains, Jamaica
Blue Mountains, Jamaica
Blue Mountains, Jamaica
Blue Mountains, Jamaica
Blue Mountains, Jamaica
Blue Mountains, Jamaica
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Continued on next page.
TREE PHYSIOLOGY ONLINE at http://heronpublishing.com
458
KNECHT AND GÖRANSSON
Table A1 Cont’d. Origin of nutrient data.
References
Species
Location
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Thompson et al. 1997
Turner 1981
Turner and Singer 1975
Turner and Singer 1975
Turner and Singer 1975
Turner et al. 1976
Turner et al. 1976
Turner et al. 1976
Turner et al. 1976
Vitousek et al. 1992
Vitousek et al. 1995
Vitousek et al. 1995
Festuca ovina
Festuca rubra
Filipendula ulmaria
Galium aparine
Galium saxatile
Geranium robertianum
Hedera helix
Helianthemum nummularium
Helictotrichon pratense
Heracleum sphondylium
Holcus lanatus
Holcus mollis
Hyacinthoides nonscripta
Juncus effusus
Juncus squarrosus
Koeleria macrantha
Lamiastrum galeobdolon
Leontodon hispidus
Lolium perenne
Lotus corniculatus
Luzula sylvatica
Mercurialis perennis
Minuartia verna
Myrrhis odorata
Nardus stricta
Origanum vulgare
Oxalis acetosella
Pilosella officinarum
Plantago lanceolata
Poa annua
Poa trivialis
Potentilla erecta
Pteridium aquilinum
Ranunculus ficaria
Rumex acetosa
Rumex acetosella
Sanguisorba minor
Scabiosa columbaria
Silene dioica
Stellaria holostea
Stellaria media
Thymus polytrichus
Tussilago farfara
Typha latifolia
Urtica dioica
Vaccinium myrtillus
Viola riviniana
Pseudotsuga menziesii
Abies amabilis (+Tsuga mertensiana)
Vaccinium deliciosum
Xerophyllum tenax
Alnus rubra
Berberis nervosa
Polystichum munitum
Pteridium aquilinum
Metrosideros polymorpha
Cheirodendron trigynum
Cibotium chammissoi
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Central England
Washington, USA
Findley Lake, Washington, USA
Findley Lake, Washington, USA
Findley Lake, Washington, USA
Allan E. Thompson Research Center, Washington, USA
Allan E. Thompson Research Center, Washington, USA
Allan E. Thompson Research Center, Washington, USA
Allan E. Thompson Research Center, Washington, USA
Hawaii, USA
Hawaii, USA
Hawaii, USA
Continued on facing page.
TREE PHYSIOLOGY VOLUME 24, 2004
NUTRIENT RATIOS IN TERRESTRIAL PLANTS
459
Table A1 Cont’d. Origin of nutrient data.
Reference
Species
Location
Vitousek et al. 1995
Vitousek et al. 1995
Vitousek et al. 1995
Vitousek et al. 1995
Vitousek et al. 1995
Vitousek et al. 1995
Vitousek et al. 1995
Vitousek et al. 1995
Whittaker et al. 1979
Whittaker et al. 1979
Whittaker et al. 1979
Whittaker et al. 1979
Whittaker et al. 1979
Woodwell et al. 1975
Woodwell et al. 1975
Woodwell et al. 1975
Woodwell et al. 1975
Woodwell et al. 1975
Woodwell et al. 1975
Woodwell et al. 1975
Woodwell et al. 1975
Foulds 1993
Foulds 1993
Foulds 1993
Foulds 1993
Foulds 1993
Foulds 1993
Foulds 1993
Foulds 1993
Foulds 1993
Foulds 1993
Foulds 1993
Foulds 1993
Foulds 1993
Foulds 1993
Foulds 1993
Foulds 1993
Foulds 1993
Foulds 1993
Foulds 1993
Foulds 1993
Foulds 1993
Foulds 1993
Foulds 1993
Foulds 1993
Foulds 1993
Foulds 1993
Foulds 1993
Foulds 1993
Foulds 1993
Foulds 1993
Foulds 1993
Foulds 1993
Foulds 1993
Foulds 1993
Foulds 1993
Foulds 1993
Foulds 1993
Cibotium glaucum
Coprosma spp.
Dicranopteris linearis
Ilex anomala
Metrosideros polymorpha, glabrous
Metrosideros polymorpha, pubescent
Myrsine lessertiana
Vaccinium calycinum
Acer saccharum
Acer spicatum
Betula lutea
Fagus grandifolia
Picea rubens
Gaylussacia baccata
Kalmis angustifolium
Pinus rigida
Quercus alba
Quercus coccinea
Quercus ilicifolia
Vaccinium angustifolium
Vaccinium vacillans
Aizoaceae
Amaranthaceae
Apiaceae
Apocynaceae
Asteraceae
Brassicaceae
Caryophyllaceae
Casuarinaceae
Chenopodiaceae
Chloantaceae
Crassulaceae
Cupressaceae
Cyperaceae
Dillaniaceae
Droseraceae
Epacridaceae
Euphorbiaceae
Frankeniaceae
Geraniaceae
Goodeniaceae
Haemodoraceae
Haloragaceae
Iridaceae
Juncaceae
Lamiaceae
Lauraceae
Leguminosae: 1. Caesalpiniaceae
2. Mimosaceae
3. Papilionaceae
Liliaceae:
1. Anthericaceae
2. Asphodelaceae
3. Colchicaceae
4. Dasypogonaceae
5. Phormiaceae
6. Xanthorrhoeaceae
Lobeliaceae
Loganiaceae
Hawaii, USA
Hawaii, USA
Hawaii, USA
Hawaii, USA
Hawaii, USA
Hawaii, USA
Hawaii, USA
Hawaii, USA
Hubbard Brook, New Hampshire, USA
Hubbard Brook, New Hampshire, USA
Hubbard Brook, New Hampshire, USA
Hubbard Brook, New Hampshire, USA
Hubbard Brook, New Hampshire, USA
Brookhaven, New York, USA
Brookhaven, New York, USA
Brookhaven, New York, USA
Brookhaven, New York, USA
Brookhaven, New York, USA
Brookhaven, New York, USA
Brookhaven, New York, USA
Brookhaven, New York, USA
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Continued on next page.
TREE PHYSIOLOGY ONLINE at http://heronpublishing.com
460
KNECHT AND GÖRANSSON
Table A1 Cont’d. Origin of nutrient data.
References
Species
Location
Foulds 1993
Foulds 1993
Foulds 1993
Foulds 1993
Foulds 1993
Foulds 1993
Foulds 1993
Foulds 1993
Foulds 1993
Foulds 1993
Foulds 1993
Foulds 1993
Foulds 1993
Foulds 1993
Foulds 1993
Foulds 1993
Foulds 1993
Foulds 1993
Foulds 1993
Foulds 1993
Foulds 1993
Foulds 1993
Foulds 1993
Loranthaceae
Myoporaceae
Myrtaceae
Orobanchaceae
Poaceae
Proteaceae
Polygolaceae
Primulaceae
Ranunculaceae
Restionaceae
Rhamnaceae
Rubiaceae
Rutaceae
Santalaceae
Sapindaceae
Solanaceae
Sterculiaceae
Stylidiaceae
Thymelaeaceae
Typhaceae
Verbeniaceae
Violaceae
Zamiaceae
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
TREE PHYSIOLOGY VOLUME 24, 2004