Potassium — A Non-Limiting Nutrient in Fresh Waters?

Author(s): J.F. Talling

Source: Freshwater Reviews, 3(2):97-104. 2010.
Published By: Freshwater Biological Association
DOI: http://dx.doi.org/10.1608/FRJ-3.2.1
URL: http://www.bioone.org/doi/full/10.1608/FRJ-3.2.1

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 97

Article

Potassium – a non-limiting nutrient in fresh waters?

J.F. Talling
Hawthorn View, The Pines, Bongate, Appleby, Cumbria CA16 6HR, UK

Received 18 November 2009; accepted 4 February 2010; published 12 July 2010

Abstract

Possible limitation of growth and distribution of freshwater organisms by the availability of potassium,
an essential major bio-constituent and plant nutrient, is considered for inland waters. It is interpreted in
›Ž•Š’˜—ȱ˜ȱ‘Žȱ›Š—Žȱ˜ȱŒ˜—ŒŽ—›Š’˜—œȱ—˜›–Š••¢ȱŽ—Œ˜ž—Ž›ŽȱŠ—ȱŽ¡™Ž›’–Ž—Š•ȱ ˜›”ȱ˜—ȱœ™ŽŒ’ęŒȱ›˜ ‘ȱ
rates of algae at low concentrations, minimum cell and biomass quotas of K, biomass yields under
graded additions, inhibition at higher concentrations, and response to cation-ratios in the medium.

The range of K+ concentrations in inland waters is surveyed. Most concentrations exceed 10 μmol L-1,
and are greatly in excess of those (under 1 μmol L-1Ǽȱ˜ž—ȱ•’–’’—ȱœ™ŽŒ’ęŒȱ›˜ ‘ȱ›ŠŽœȱ˜ȱŽœȱœ™ŽŒ’Žœǯȱȱ
They are also in excess of the content in most natural populations of phytoplankton when yields have
the experimental minimum or limiting cell/biomass quota, of the order of 1 % dry weight. Limiting
Œ˜—ŒŽ—›Š’˜—œȱ˜›ȱœ™ŽŒ’ęŒȱ›˜ ‘ȱ›ŠŽȱ’—ȱ—Šž›ŽȱŠ›Žȱ‘Ž›Ž˜›Žȱ™›˜‹Š‹•¢ȱ›Š›Žǯȱȱ‘Ž¢ǰȱŠ—ȱ¢’Ž•ȱ•’–’Š’˜—œǰȱ
–’‘ȱ™˜œœ’‹•¢ȱ‹Žȱ›ŽŠŒ‘ŽȱŠĞŽ›ȱŽ¡›Ž–ŽȱŽ™•Ž’˜—ȱ‹¢ȱŽ—œŽȱœŠ—œȱ˜ȱŠšžŠ’Œȱ–ŠŒ›˜™‘¢ŽœDzȱœ˜–Žȱ
depletions are recorded for a charophyte and water-cress (Rorippa nasturtium-aquaticum). Retention
‹¢ȱœŽŠœ˜—Š•ȱŸŽŽŠ’˜—ȱ’—ȱ‘ŽȱŒŠŒ‘–Ž—ȱŒŠ—ȱ–˜›Žȱ‘Š—ȱ˜ěœŽȱŽ•’ŸŽ›¢ȱ’—ȱ›Š’—Š••ȱŠ—ȱ›Žœž•ȱ’—ȱ–’—’–Š•ȱ
concentrations of between 1 μmol L-1 and 5 μmol L-1. There is evidence that these concentrations,
combined with poor nutrition, can limit the distribution of larger Crustacea in some upland streams.
Other less substantiated sources of limitation in nature relate to ratios of cationic concentrations,
’—ȱ™Š›ȱ˜›ȱž—Œ’˜—ȱ˜ȱŠ•Š•ȱ̊Ž••Šǯȱȱ‘Ž›Žȱ’œȱœ˜–ŽȱŽ¡™Ž›’–Ž—Š•ȱŽŸ’Ž—ŒŽȱ˜›ȱ›˜ ‘ȬȱŠ—ȱ¢’Ž•Ȭ
limitation of species of the chrysophyte Dinobryon by higher and naturally occurring K+ concentrations.

Keywords: bioassay; Crustacea; growth rate; inhibition; macrophytes; nutrient depletion; phytoplankton;
potassium; yield.

Introduction planktonic micro-vegetation and its production in inland

waters. Summaries of this experience and texts of general
Potassium is well known as a constituent of common freshwater science (Hutchinson, 1957, 1967; Wetzel, 2001;
NPK fertilisers and as a potential or actual limiting Ž¢—˜•œǰȱŘŖŖŜǼȱŽ—Ž›Š••¢ȱ’ŸŽȱ•’Ĵ•Žȱœ™ŠŒŽȱ˜ȱ™˜Šœœ’ž–ǯȱȱ ȱ
factor for the growth of terrestrial vegetation. A very appears as a ubiquitous major cation, in quantities likely to
’쎛Ž—ȱ ’–™›Žœœ’˜—ȱ ‘Šœȱ Ž–Ž›Žȱ ›˜–ȱ Ž¡™Ž›’Ž—ŒŽȱ ’‘ȱ exceed those involved in planktonic uptake and to escape

DOI: 10.1608/FRJ-3.2.1 Freshwater Reviews (2010) 3, pp. 97-104

© Freshwater Biological Association 2010
98 Talling, J.F.

conspicuous depletions such as are commonplace for Concentration in inland waters

phosphorus, inorganic nitrogen and silicate-silicon.
All this is broadly true, but …. The following review Potassium is one of four cations – Ca2+, Mg2+, Na+ and K+
takes up reasons for renewed interest by the freshwater – that collectively account for most of the total cationic
ecologist in this physiologically unique element. concentration in most natural waters (very acid waters,
with more H+, excepted). It is usually the least abundant
Potassium as a bio-component of these four major cations as assessed, for example, in
–’Œ›˜–˜•Žœȱ ǻΐ–˜•Ǽȱ ™Ž›ȱ ǰȱ Šȱ œŒŠ•Žȱ ‘Ž›ŽŠĞŽ›ȱ Ž–™•˜¢Žȱ ǻŗȱ
There follows a brief reminder of some salient facts. μmol = 1 μequiv = 0.0391 mg). However, local exceptions
Potassium is the predominant cation associated with exist, as for instance in drainage from the Alsace potash
cytoplasm in the cytosol. In plant material it is present sources to the middle Rhine that, due to accompanying
there in considerable concentrations of the order of 40 sodium chloride, has introduced episodes of toxicity to
-1
mmol L (Raven, 1984) and is the usual major cation in growers of tomatoes in the Netherlands (Golterman, 1975).
the more variable contents of the vacuole (Leigh & Wyn Contrasts between K+ and Na+ regarding abundance
Jones, 1986). Hence it is osmotically active, i.e. an osmolyte. and behaviour are evident in both geochemistry and
Aquatic macrophytes typically contain the element in the physiology, despite much chemical similarity and
›Ž’˜—ȱ ˜ȱ Řȱ Ɩȱ ˜ȱ Śȱ Ɩȱ ˜ȱ ‘Žȱ ›¢ȱ Ž’‘ǰȱ ˜ĞŽ—ȱ œ’–’•Š›ȱ ’—ȱ near-equal primary abundance in the earth’s crust. In
magnitude to the content of nitrogen (examples in Talling seawater the absolute, if not relative, concentration of K+
ǭȱŽ–˜Š••ŽǰȱŗşşŞǰȱ’ǯȱŘǯřŚǼǯȱȱ œȱž™Š”Žȱ’œȱ’—ĚžŽ—ŒŽȱ‹¢ȱ is considerable, around 9700 μmol L-1 for normal salinity.
electrical properties of the cell membrane, and involves At another extreme, it has been recorded within the
one or more ion-pumps that may be sensitive to light and concentration range 1 μmol L-1 to 50 μmol L-1 in mostly
to other cations (e.g. Barber, 1968; Shiel & Barber, 1971). low-solute fresh waters resulting from drainage over
+
There are non-reciprocal relations with NH transport.4
impervious rocks. Examples in Africa and Amazonia are
Ž•Š’˜—œȱ ’‘ȱ–Ž–‹›Š—Žȱ™Ž›–ŽŠ‹’•’¢ȱŒŠ—ȱ‹Žȱ’—ĚžŽ—ŒŽȱ instanced by Beadle (1981) and others in the Pyrenees and
by the ratio of monovalent to divalent cations in the Alps by Psenner & Catalan (1994). Such waters are usually
Ž¡Ž›—Š•ȱ–Ž’ž–ǯȱȱ —ȱ̊Ž••ŠŽœǰȱŠ—ȱ’—ĚžŽ—ŒŽȱ˜—ȱ̊Ž••Š›ȱ ŽŠœ’•¢ȱ’œ’—ž’œ‘ŽǰȱŽŸŽ—ȱ’—ȱ‘Žȱꎕǰȱ‹¢ȱ‘Ž’›ȱ•˜ ȱŽ•ŽŒ›’ŒŠ•ȱ
activity also involves cation concentrations and ratios (e.g. conductivity. It is likely that many K+ concentrations
Nultsch, 1979). determined before about 1965 are over-estimates, due to
Potassium is a co-factor in many enzymic reactions. Its ’ĜŒž•’Žœȱ ˜ȱ ž••¢ȱ Œ˜–™Ž—œŠ’—ȱ ˜›ȱ ’—Ž›Ž›Ž—ŒŽȱ ›˜–ȱ
exchanges with the outer medium may involve biological ˜‘Ž›ȱŒŠ’˜—œȱ ’‘ȱ‘Žȱ̊–Žȱ™‘˜˜–Ž›’ŒȱǻŠ˜–’ŒȱŽ–’œœ’˜—Ǽȱ
›‘¢‘–œǰȱ Šœȱ ’—ȱ ‘Žȱ ’—˜ĚŠŽ••ŠŽȱ Gonyaulax polyedra method then most used. Later values obtained by atomic
ǻ ŽŽ—Ž¢ǰȱ ŗşŝŚǼǰȱ Š—ȱ ‹Žȱ ’—ĚžŽ—ŒŽȱ ‹¢ȱ Š+ as an uptake absorption spectrophotometry had less exposure to
co-substrate (Maathius et al., 1996). In aquatic animals interference and greater sensitivity, with a limit of detection
as well as plants, interrelations between internal and by normal procedure (without pre-concentration) about
external concentrations are governed by two-way dynamic 0.3 μmol L-1. These include K+ concentrations as low as 10
exchange, complemented by ingested quantities in μmol L-1 in water with considerable total solutes derived
phagotrophs. Exchange is accentuated when relatively from drainage in karst landscapes. The upland Malham
impervious barriers (e.g. exoskeletons) are absent, resulting Tarn in Yorkshire is one example, to which we will return
’—ȱ Šȱ ›Š™’ȱ ž›—˜ŸŽ›ȱ ‘Šȱ –Š¢ȱ ‘ŠŸŽȱ œ’—’ęŒŠ—ȱ Ž—Ž›¢ȱ later. In the English Lakes, of low solute concentration,
demand. K+ is one of several major ionic constituents positively
Œ˜››Ž•ŠŽȱ ’‘ȱ ‹’˜•˜’ŒŠ•ȱ ™›˜žŒ’Ÿ’¢ǯȱ ȱ ‘žœȱ žŒ•’ěŽȱ Žȱ
al. (1982) distinguished three groups of these lakes with

© Freshwater Biological Association 2010 DOI: 10.1608/FRJ-3.2.1

Potassium – a non-limiting nutrient in fresh waters? 99

increasing abundance of planktonic algae, in which mean ‘Žȱ ’—œŠ—Š—Ž˜žœȱ ˜›ȱ œ™ŽŒ’ęŒȱ ›˜ ‘ȱ ›ŠŽǰȱ ›Š‘Ž›ȱ ‘Š—ȱ
K+ concentrations were 7–11, 9–15 and 17–26 μmol L-1. ’—Œ›ŽŠœŽȱ˜ȱ¢’Ž•ȱŠĞŽ›ȱŠ—ȱŠ›‹’›Š›¢ȱ’–Žȱ™Ž›’˜ǯȱȱ —ȱœŽ–’Ȭ
Three groups of upland tarns in the same district averaged logarithmic plots of population density against time, with
5–7 μmol L-1. ‹ŠŒ‘ȱŒž•ž›Žœǰȱ‘Žȱ›Š’Ž—ȱ˜ȱŠȱ™•˜ȱ’œȱŠȱ–ŽŠœž›Žȱ˜ȱœ™ŽŒ’ęŒȱ
+
The highest K concentrations are most likely to be growth rate, in ln units per unit time.
found in those inland waters within closed drainage Such a plot is illustrated in Fig. 1, from Jaworski et al.
basins, lacking an outlet, that are subject to evaporative (2003), which shows growth of the common planktonic
concentration in arid climates. There are numerous diatom Asterionella formosa in culture media that initially
examples in Africa, for which Talling (1992) lists records contained the low K+ concentrations of 0.7, 1.3 and 6.0
+ -1 + -1
in the range 500 μmol K L to 10 000 μmol K L . μmol L-1ǯȱȱ••ȱŒž•ž›Žœȱ›Ž ȱŠȱŸŽ›¢ȱœ’–’•Š›ȱœ™ŽŒ’ęŒȱ›ŠŽœȱ
+
Because of its considerable concentration in sea- ˜›ȱ ‘Žȱ ꛜȱ śȱ Š¢œǰȱ ‹žȱ ‘Ž›ŽŠĞŽ›ȱ ‘Žȱ •Š›Ž›ȱ ’—’’Š•ȱ
ŠŽ›ǰȱ™˜Šœœ’ž–ȱ’œȱœ’—’ęŒŠ—ȱŠ–˜—ȱ‘ŽȱȁŒ¢Œ•’Œȱ
salts’ derived from sea-spray and carried in
rain to inland waters. This source accounts for
some enhancement of the K+ concentrations of
inland waters in oceanic regions, increasing with
proximity to the sea. A further but more secondary
source is from the periodic decomposition of
˜›Š—’Œȱ –ŠĴŽ›ǰȱ œžŒ‘ȱ Šœȱ ‘Žȱ Šžž–—Š•ȱ ’—™žȱ ˜ȱ
leaves and dead herbaceous vegetation. Especially
in some upland and glaciated landscapes, the
K+ concentrations in stream water can show an
autumnal pulse but later decrease to low values –
e.g. 1–5 μmol L-1 – in mid-summer from retention
’—ȱ‘ŽȱŒŠŒ‘–Ž—Dzȱ‘ŽȱŽŒ›ŽŠœŽȱ’œȱ’››Žž•Š›•¢ȱ˜ěœŽȱ
by K+ input in rainwater. Examples from long-
term records in the English Lake District are
ŽœŒ›’‹Žȱ‹¢ȱžŒ•’ěŽȱǭȱŠ››’Œ”ȱǻŗşŞřǼǯȱȱ‹ž—Š—ȱ
›ŽŽȬœ Š–™ȱ–Š¢ȱŠŒȱŠœȱŠȱ›˜œœȱȁ—ž›’Ž—ȱ™ž–™Ȃǰȱ
although unlike one active across membranes
(cf. Denny, 1985), the major sites of uptake and
release by decay being vertically separated.

Limiting concentrations

Aware of the wide range of K+ concentrations in

inland waters, we may ask where, if anywhere,
it includes waters limiting for growth of free-
Fig. 1. Growth of the plankton diatom Asterionella formosa in culture media A–C
̘Š’—ȱŠž˜›˜™‘œȱœžŒ‘ȱŠœȱ™‘¢˜™•Š—”Ž›œǯȱȱ Ž›Žȱ with three initial concentrations of K+, including supplementation at day 7 with
experimental work with cultures is required. CO2 (A1–C1), broken lines. Below, corresponding sequential measurements of
total K+ and dissolved K+ (broken lines), with divergence because of increase
Growth is most fundamentally measured by that in the particulate K of algal biomass. From Jaworski et al. (2003, Fig. 1), with
characteristic of exponential replicative increase, correction of cell concentration units.

DOI: 10.1608/FRJ-3.2.1 Freshwater Reviews (2010) 3, pp. 97-104

100 Talling, J.F.

Œ˜—ŒŽ—›Š’˜—œȱ ›Žœž•Žȱ ’—ȱ ™›˜™˜›’˜—ŠŽ•¢ȱ ›ŽŠŽ›ȱ ꗊ•ȱ the challenge of K+ concentrations less than 10 μmol L-1 in
yields – that could be further increased slightly by CO2 situations of poor-quality food. This could account for the
enrichment. Direct measurements of the dissolved and absence of G. pulex from most of the upper River Duddon
total (dissolved + particulate) K+ showed decline of the tributaries, where the diet available is less suitable and
former during growth. These results were evidence that such low concentrations of K+ are frequent during summer
+
growth rate was independent of K concentration down due to accumulation in vegetation within the catchment.
-1
to 0.7 μmol L , not far removed from the analytical limit
of detection (0.3 μmol L-1Ǽǯȱ ȱ ˜ ŽŸŽ›ǰȱ ‘Žȱ ꗊ•ȱ ›˜ ‘ȱ Transient depletions by plant
+
yields were dependent on K concentration in the range growth
used, consistent with a minimum cell quota of 2.8 μmol
per 108 cells, approximately 1 % of the organic dry weight Figure 1 illustrates how the growth of phytoplankton can
Ȯȱ Šȱ ŸŠ•žŽȱ ’‘’—ȱ ‘Žȱ ›Š—Žȱ ˜ž—ȱ ‹¢ȱ Ž›•˜ěȱ ǭȱ ’œ‘‹ŽŒ”ȱ deplete the concentration of K+ in the medium, and how
(1969) to be limiting for other algae. The yield dependence the extent of depletion increases with the magnitude of
may partly account for varying older estimates of K+ population increase. Clearly the relative depletion will
limitation of phytoplankton at higher concentrations depend on the ratio between the algal crop-potassium and
in nature (Höll, 1951) and culture (MacCarthy & the ambient K+ȱ’—’’Š••¢ȱ™›ŽœŽ—ȱ’—ȱž—’ȱŸ˜•ž–Žǯȱȱ ȱ‘Žȱ•ŠĴŽ›ȱ
ŠĴŽ›œ˜—ǰȱ ŗşŝŚDzȱ ŽŽ Š•ȱ Žȱ Š•ǯǰȱ ŗşŞŗǼǯȱ ȱ ŽŸŽ›‘Ž•Žœœǰȱ is large, the relative depletion is likely to be minor and
‘Žȱ ž—Š–Ž—Š•ȱ ꗍ’—ȱ ‘Šȱ œ™ŽŒ’ęŒȱ ›˜ ‘ȱ ›ŠŽȱ Šœȱ ’ĜŒž•ȱ˜ȱŽŽŒȱȮȱž—•ŽœœȱŸŽ›¢ȱ•Š›Žȱ’—Œ›Ž–Ž—œȱ˜ȱ‹’˜–Šœœȱ
+ -1
independent of low K concentrations down to 1 μmol L are involved. If the content of algal K is of the order of 1 %
or less, shown to be applicable to two further species, of the organic dry weight, it can be deduced that algal crops
is ecologically most relevant. It would account for the in the common range of 1–10 mg dry organic weight per
general failure to identify potassium as a limiting element L would remove less than 13 % of the K+ content in those
in fresh waters by bioassay additions (e.g. Rodhe, 1948). most frequent natural concentrations above 20 μmol L- 1.
There appears to be an interesting parallel situation Such minor depletion is in accord with the pragmatic
with some aquatic animals, such as freshwater conclusion that appreciable K+ decline rarely accompanies
+
Malacostraca (Crustacea), that have a minimum K increase in natural micro-algal populations (e.g. Wetzel,
requirement (e.g. Shaw, 1959). This was considered by ŘŖŖŗǼǰȱŠ•‘˜ž‘ȱ’ȱ–žœȱ‹ŽȱŠ–’ĴŽȱ‘Šȱ‘Ž›Žȱ‘ŠŸŽȱ‹ŽŽ—ȱ
Beadle (1981) as involving concentrations too low, relative Ž ȱœ¢œŽ–Š’ŒȱŠĴŽ–™œȱ˜ȱ˜••˜ ȱœžŒ‘ȱœŽšžŽ—ŒŽœǯ
to natural concentrations, to be of widespread ecological Much greater increase of biomass per unit volume
œ’—’ęŒŠ—ŒŽȱ’—ȱ›’ŒŠ—ȱ›Žœ‘ȱ ŠŽ›œȱȮȱŠ•‘˜ž‘ȱŽ¡ŒŽ™’˜—œȱ is found with some stands of aquatic macrophytes. As
existed, as at Lake Nabugabo near Lake Victoria. Š•›ŽŠ¢ȱ—˜Žǰȱ‘ŽœŽȱ˜ĞŽ—ȱ‘ŠŸŽȱŠȱ›Ž•Š’ŸŽȱ ȱŒ˜—Ž—ȱ–žŒ‘ȱ
’••˜ž‘‹¢ȱǭȱžŒ•’ěŽȱǻŗşŝŜǼȱŠ—ȱžŒ•’ěŽȱǭȱŠ››’Œ”ȱ greater than 1 % of dry weight. They include charophytes,
(1983) have assessed more extensive information from ’‘ȱȁ’Š—ȱŒŽ••œȂȱ˜›ȱŒ˜Ž—˜Œ¢Žœȱ ‘’Œ‘ȱ‘ŠŸŽȱ‹ŽŽ—ȱŠŸ˜ž›’Žȱ
English fresh waters in relation to their own and other subjects for experimental studies of ion uptake. There is
experimental work on the larger Crustacea. It appears that documentation (Talling & Parker, 2002b) of the K+ relations
starved mature inter-moult individuals of the common of a stand of Chara globularis var. virgata in Malham Tarn,
freshwater shrimp Gammarus pulex can only maintain Yorkshire, where – as already mentioned – the ambient
minimum levels of potassium balance for short periods K+ȱŒ˜—ŒŽ—›Š’˜—ȱ’œȱ•˜ ǯȱȱ‘Žȱ•ŠĴŽ›ȱŒ˜—’’˜—ȱ’œȱŠ™™Š›Ž—•¢ȱ
(days) at an external concentration of 10 μmol L-1 to not uncommon in Chara lakes (e.g. McConnaughey et al.,
15 μmol L-1, whereas well-fed individuals survived for ŗşşŚǼǯȱȱ˜–™Š›’œ˜—ȱ‘Šœȱ‹ŽŽ—ȱ–ŠŽȱ˜ȱ’—Ě˜ ȱŠ—ȱ˜žĚ˜ ȱ
-1
much longer periods at 5 μmol L . Immature animals which concentrations. The results (Fig. 2) show a divergence
need to grow and moult at frequent intervals may not meet indicative of loss in transit, that corresponds to the main

© Freshwater Biological Association 2010 DOI: 10.1608/FRJ-3.2.1

Potassium – a non-limiting nutrient in fresh waters? 101

growth season of the Chara. It is consistent with the reported of under 1.3 μmol L-1 approximate the levels
measured K content of biomass and the estimated biomass at which the direct evidence was found for growth
+
increment, allowing for some uptake of K via rhizoids •’–’Š’˜—ȱ ˜ȱ œ™ŽŒ’ęŒȱ ›˜ ‘ȱ ›ŠŽȱ ‹¢ȱ ™‘¢˜™•Š—”˜—ǯ
from the sediment. The range of seasonal K+ depletion Experimental studies have also been undertaken to
-1
is from c. 10 to 1 μmol L ǯȱȱ‘Žȱ•ŠĴŽ›ȱ–’—’–ž–ȱ’œȱŠ‹˜ŸŽǰȱ ŽŽŒȱ ž™Š”Žȱ ˜ȱ —ž›’Ž—œȱ ž›’—ȱ œ›ŽŠ–ȱ ̘ ǰȱ žœ’—ȱ
but not greatly above, the markedly sub-micromolar successive stations below an input of mixed nutrients
concentrations that were limiting to growth of the planktonic accompanied by a conservative tracer. Ratios of
Asterionella formosa (Fig. 1). Other studies (Proctor, 1995) nutrient to tracer can then be used to gauge nutrient
have been made of K+ concentrations in water from areas ž™Š”Žȱ ’—ȱ ›Ž•Š’˜—ȱ ˜ȱ ’–Žȱ Š—ȱ ’œŠ—ŒŽȱ ˜ȱ ̘ ȱ ǻŒ˜••ǰȱ
of raised bog and fen beside the Tarn, that indicated a 1974; Stream Solute Workshop, 1990). Comparative
rapid turnover. A depletion during summer was detected tests with salts yielding potassium, ammonium and
in an American Chara lake (McConnaughey et al., 1994). phosphate ions in an acid upland stream demonstrated
Another input–output situation that interposes a progressive uptake of ammonium and phosphate,
large macrophyte biomass concerns the beds of water- but not of potassium (Talling & Parker, 2002a).
cress, Rorippa nasturtium-aquaticum (L.) Hayek (Nasturtium
˜ĜŒ’—Š•Ž Brown), that are present in the headwaters of Limitation by K+ toxicity
several chalk streams in southern England. Here there
has been a commercial practice of adding K-containing Besides the potential for growth limitation at low
fertiliser to increase yields of water-cress. Several concentrations, potassium and many other ions can be
studies (Crisp, 1970; Casey, 1981; Casey & Smith, 1993) ˜¡’ŒȱŠȱœžĜŒ’Ž—•¢ȱ‘’‘ȱŒ˜—ŒŽ—›Š’˜—œǯȱȱ‘’œȱ‘ŠœȱŽŒ˜•˜’ŒŠ•ȱ
+
have demonstrated a strong seasonal uptake of K with relevance if the range for toxicity overlaps the range
Ž™•Ž’˜—ȱ ‹Ž ŽŽ—ȱ —Šž›Š•ȱ ’—Ě˜ ȱ ǻŽǯǯȱ ˜ȱ ›˜ž— ŠŽ›ǰȱ commonplace in natural waters. Following some early
-1
mean 22 μmol L Ǽȱ Š—ȱ ˜žĚ˜ ǯȱ ȱ ‘Žȱ Ž¡›Ž–Žȱ –’—’–Šȱ experimental work by Rodhe (1948), phosphate was widely
supposed (e.g. by Hutchinson, 1967) to be
in this category for species of freshwater
Œ‘›¢œ˜ĚŠŽ••ŠŽœȱ œžŒ‘ȱ Šœȱ Dinobryon spp.
Further support came from supposedly
Œ˜››Ž•Š’ŸŽȱ ꎕȱ ŠŠȱ ˜—ȱ Œ‘›¢œ˜™‘¢Žȱ
seasonality, including that of Pearsall
(1932). However, others – including
myself (Talling, 1962) – later found that
Dinobryon could be cultured in phosphate-
rich media. As K+ was the balancing cation
in the experimental enrichments of Rodhe,
Lehman (1976) was led to test whether
it rather than phosphate was a growth-
inhibiting agent at ecologically relevant
concentrations. He found that the growth
yield of several species of Dinobryon was
markedly reduced in cultures when K+
Fig. 2. Malham Tarn, Yorkshire: (a) timing of seasonal growth of Chara globularis, -1
indicated by the mean length of shoots during 1980; (b) potassium uptake, indicated by concentrations exceeded 100 μmol L ,
‘Žȱ’쎛Ž—ŒŽȱ˜ȱŒ˜—ŒŽ—›Š’˜—œȱ’—ȱ’—Ě˜ ȱŠ—ȱ˜žĚ˜ ȱ ŠŽ›ȱž›’—ȱŗşŞŝǯȱȱ˜’ꮍȱŠĞŽ›ȱ and possibly would occur as low as
Pentecost (1984) and Talling & Parker (2002a).

DOI: 10.1608/FRJ-3.2.1 Freshwater Reviews (2010) 3, pp. 97-104

102 Talling, J.F.

25 μmol L-1 under some conditions that included paucity species are likely to be limited or greatly reduced – in
of Ca2+ and Mg2+. Sandgren (1988) gives further support. the absence of replenishment – by crop yields of even
+
In contrast, some classical culture media contain K (e.g. as exceptionally dense natural populations.
KNO3) at much higher concentrations, but were not widely Nevertheless, there are reasons for not entirely
successful for phytoplankton. The major advance in the eliminating the possibilities of K+ limitation or toxicity.
culture of freshwater phytoplankton came when Chu Instances of ambient K+ depletion by aquatic macrophytes
(1942) successfully tested a range of more dilute media could proceed further than in the examples cited, to sub-
based on analogy with natural lake waters. We have seen, micromolar levels not much lower than the concentration
+ -1
however, that K concentrations of 100 μmol L or more reached in Malham Tarn. If achieved, such levels are likely
are found in a range of inland waters. One major group, ˜ȱ•’–’ȱ‹˜‘ȱœ™ŽŒ’ęŒȱ›˜ ‘ȱ›ŠŽœȱŠ—ȱ™˜Ž—’Š•ȱ‹’˜–Šœœȱ
documented by Talling & Talling (1965) and inserted in yield of phytoplankton. Estimates of sensitivity to depletion
+
Fig. 3ǰȱ’œȱ’—ĚžŽ—ŒŽȱ‹¢ȱ -rich drainage from K-rich lavas based on a cell or biomass quota have various uncertainties,
˜ȱ‘ŽȱŽœȱ’Ğȱ’—ȱ›’ŒŠǯȱȱ ȱ’—Œ•žŽœȱ•Š”Žœȱ•‹Ž›ǰȱ Š›ǰȱ including possible seasonal variation of K content
Kivu and Tanganyika. (described for a Chara by Winter et al., 1987), and sensitivity
Another origin of unfavourable aqueous environments of the ion-uptake pump to light and the concentrations of
+
’—ĚžŽ—ŒŽȱ ‹¢ȱ concentration might exist if cationic other ambient cations. Finally, if the evidence that higher
›Š’˜œȱ Š›Žȱ Šȱ ŠŒ˜›ȱ ’—ĚžŽ—Œ’—ȱ ›˜ ‘ǯȱ ȱ ‘’œȱ ™˜œœ’‹’•’¢ȱ natural K+ȱ Œ˜—ŒŽ—›Š’˜—œȱ ŒŠ—ȱ ‹Žȱ ˜¡’Œȱ ’œȱ Œ˜—ę›–Žȱ Š—ȱ
was raised by Pearsall (1930, 1932) from correlations in his extended, it might reopen the question of K+ȱ ’—ĚžŽ—ŒŽȱ
pioneer ecological work on the English Lakes. Subsequent Ÿ’Šȱ Š–‹’Ž—ȱ ŒŠ’˜—ȱ ™›˜™˜›’˜—œǰȱ Šȱ •ŽŠœȱ ˜›ȱ ̊Ž••ŠŽœǯ
experimental tests and evidence from distribution in Limitation or restriction by potassium should not
—Šž›Žȱ ǻž—ǰȱ ŗşŜśDzȱ ˜œœǰȱ ŗşŝŘȮŝřǼǰȱ ‘ŠŸŽȱ ˜ěŽ›Žȱ •’Ĵ•Žȱ always be neglected.
support to this hypothesis. It has, however, been raised
by Wood & Talling (1988) in connection with a supposed Acknowledgements
correlation between the ratio Ca2+ + Mg2+ / Na+ + K+ and the
˜ŒŒž››Ž—ŒŽȱ ˜ȱ ̊Ž••ŠŽȱ ™‘¢˜™•Š—”Ž›œȱ ’—ȱ ›’ŒŠ—ȱ •Š”Žœǯ This review was stimulated by experimental work with
George Jaworski and Ivan Heaney on the limitation by
Concluding remarks ™˜Šœœ’ž–ȱ˜ȱŠ•Š•ȱ›˜ ‘ȱ›ŠŽȱŠ—ȱ¢’Ž•ǯȱȱ ȱ‘Šœȱ‹Ž—ŽęŽȱ
›˜–ȱ‘Ž•™ž•ȱŒ˜––Ž—œȱ‹¢ȱ ˜‘—ȱŠŸŽ—ȱǰȱŠŸ’ȱžŒ•’ěŽȱ
The foregoing survey can be summarised in relation and two anonymous reviewers. John Wiley & Sons kindly
to a scale of K+ concentration in inland waters (Fig. 3). gave permission for the reproduction of material in Figures
Concentrations encountered are normally greater than 1 and 3, originally published in the journal Freshwater
‘˜œŽȱ ‹¢ȱ ‘’Œ‘ȱ œ™ŽŒ’ęŒȱ ›˜ ‘ȱ ›ŠŽœȱ ˜ȱ ™‘¢˜™•Š—”˜—ȱ Biology.

Fig. 3. Logarithmic scale for concentration of potassium, with inserted examples from various natural waters and culture media; also limit
˜ȱŠ—Š•¢’ŒŠ•ȱŽŽŒ’˜—ȱ‹¢ȱŠ˜–’ŒȱŠ‹œ˜›™’˜—ȱœ™ŽŒ›˜œŒ˜™¢ǯȱȱ˜’ꮍȱ›˜–ȱ Š ˜›œ”’ȱŽȱŠ•ǯȱǻŘŖŖřǰȱ’ǯȱř‹Ǽǯ

© Freshwater Biological Association 2010 DOI: 10.1608/FRJ-3.2.1

Potassium – a non-limiting nutrient in fresh waters? 103

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Jack Talling has been engaged, since 1950, in studies
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on the physiological ecology of freshwater algae
polyedra and phase changes in the circadian rhythm of
and the associated physical and chemical behaviour
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limnology developed from early experience in Africa
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50 years with the Freshwater Biological Association
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at Windermere, including a late period there as an
lakes and wetlands. Revue d’Hydrobiologie tropicale 25, 87-144.
Honorary Research Fellow.

© Freshwater Biological Association 2010 DOI: 10.1608/FRJ-3.2.1