154 Potassium nutrition in grapevines Australian Journal of Grape and Wine Research 9, 154–168, 2003

A review of potassium nutrition in grapevines

with special emphasis on berry accumulation
BUSSAKORN S. MPELASOKA1,4, DANIEL P. SCHACHTMAN1,3, MICHAEL T. TREEBY2
and MARK R. THOMAS1
1
CSIRO Plant Industry, Horticulture Unit, GPO Box 350 , Glen Osmond, SA 5064, Australia
2
CSIRO Plant Industry, Horticulture Unit, Private Mail Bag, Merbein,Vic. 3505, Australia
3
Present address: Donald Danforth Plant Science Center, 975 N. Warson Road, St Louis, MO 63132
4
Corresponding author: Dr Bussakorn Mpelasoka, facsimile +61 8 8303 8601, email Bussy.Mpelasoka@csiro.au

Abstract
Potassium (K) is essential for vine growth and yield. Grape berries are a strong sink for K, particularly
during ripening. Excess K levels in grape berries may have a negative impact on wine quality, mainly
because it decreases free tartaric acid resulting in an increase in the pH of grape juice, must and wine. In
Australia, high K status is common in most vineyards, which reflects the high K and high pH values of
most Australian grape juice. This necessitates pH adjustment during the vinification process, and tartaric
acid addition is a common practice in most Australian wineries. High K concentration may also lead to
excessive loss of the additional tartaric acid by precipitation as potassium bitartrate and, as a consequence,
pH adjustment becomes more difficult and expensive. Ensuring naturally low K levels in the berry will
help reduce costs of input and waste management at the winery. Potential vineyard management options
to manipulate berry K accumulation include selective use of rootstock/scion combination, canopy
management and irrigation strategies. However, the impact of these practices on determining the
optimum K concentration requires careful calibration of production parameters and the desirable grape
juice and wine quality in relation to tissue K concentration. This paper reviews and discusses the possible
functions of K in grape berries, translocation of K into the berry, and genetic and cultural factors that may
affect the accumulation of K in the berry. This will help to identify the key research and management
strategies needed for controlling K concentrations in grape berries.

Keywords: grapevine, Vitis vinifera L., potassium nutrition, K+ transport, berry potassium, wine quality

1. Potassium and wine quality
Factors influencing the pH of grape juice include the
stoichiometric exchange of organic acid hydrogen ions
with cations, leading to a decrease in free acid, and the
tartrate:malate ratio (Gawel et al. 2000). Potassium is the
major cation in grape juice. High juice K decreases free
acids and increases overall pH. Tartaric acid is a signifi-
cantly stronger acid than malic acid, therefore, at similar
values of total acidity, a lower tartrate:malate ratio may
result in a less acidic pH (Boulton 1980, Gawel et al.
2000). High malate also enhances malolactic fermenta-
tion, a secondary fermentation carried out by many lactic
acid bacteria and which may have either positive or neg-
ative impacts on the organoleptic quality of the wines.
This necessitates the use of commercial lactic acid bacteria
starter-cultures to control malolactic fermentation.
Tartrate also gives a crisp and fresh acid taste to the wine
(Rühl 2000) and is therefore more preferred than malate.
In a berry, high K may decrease the rate of malate degra-
dation through malate respiration by impeding malate
transfer from the vacuole storage pools to the cytoplasm,
the site of malate degradation (Hale 1977). During wine-
making, high K increases the precipitation of tartrate in
salt form and hence decreases free tartrate. Therefore
high K can lead to reduced tartrate:malate ratio which is
undesirable for high quality wines.
Grape juice with a high pH often results in unstable
musts and wines that are more susceptible to oxidative
and biological spoilage, and often produces a wine with
high pH and low acidity with a flat taste (Somers 1977).
The high pH of grape juice and wine also decreases the
colour quality of red wines. The degree of ionisation of
anthocyanins, which is the percentage of total antho-
cyanins present in the coloured forms, decreases as pH
increases (Somers 1975). Anthocyanins are located in
berry skin (Somers and Pocock 1986) where K concen-
tration is generally higher than in the pulp (Iland and
Coombe 1988, Walker et al. 1998). Therefore, berry K
levels are often a more important consideration for red
wine than for white wine because during the fermenta-
tion of red wine, the skin is left for some period after
crushing for the extraction of anthocyanins, during
which time more K may also be extracted. While the
optimum pH range for red wines is generally considered
to be 3.3–3.7, many Australian wines in the early years of
Australian winemaking had pH values above 3.7. These
Mpelasoka, Schachtman,Treeby & Thomas Potassium nutrition in grapevines 155

Figure 1. This model shows

the pattern of K accumulation
by grape berries in relation to
the pattern of grape berry
growth and development. Also
shown are the proposed
relative proportions of the
inflow of xylem and phloem
saps into the berry. The
Berry size
relative proportions before
veraison is not clearly known,
but here illustrated to be

Berry K content (mg/berry)

higher for xylem than phloem.
Berry size

After veraison, the inflow of

xylem sap decreases or
Berry K content
ceases, which may depend on
varieties, while the inflow of
phloem sap increases until
near harvest when it
decreases. Diagram of grape
berry growth and development
was taken from Coombe
(2001) with slight
modification. Data on berry K
accumulation were taken from
Ollat and Gaudillère (1996).

higher pH values are associated with high K levels.
Investigation of dry red wines from 75 vintages in all
Australian wine districts indicated high pH (3.7–4.3) and
high K levels (27–71 mmol/L) (Somers 1975). Reported
average values of K levels for foreign red wines are
much lower – for example, 22–32 mmol/L for the
Bordeaux reds (Somers 1977).
Although pH is commonly adjusted in Australia
during the vinification process by the addition of tartaric
acid, part of the additional tartaric acid may be precipi-
tated as potassium bitartrate due to the high K concen-
tration. This is because solubility of potassium bitartrate is
very low and the solubility decreases as K concentration
and/or pH increases. Therefore, ensuring naturally low K
levels in the berry is very desirable because it would
reduce input and waste management costs at the winery.
2. Functions of K in plants and possible roles of K
in grape berries
Potassium is an essential element for all living organisms.
The important roles of K in plants can be grouped into
four physiological-biochemical roles: (1) enzyme activa-
tion (Leigh and Wyn Jones 1984, Walker et al. 1998); (2)
cellular membrane transport processes and translocation
of assimilates (Salisbury and Ross 1992, p. 157, Patrick et
al. 2001); (3) anion neutralisation, which is essential in
maintenance of membrane potential (Maathuis and
Sanders 1996, Leigh 2001); and (4) osmotic potential
regulation, which is one of the important mechanisms in
the control of plant water relations (Davies and Zhang
1991), turgor maintenance and growth. Although other
cations may replace K in some of these functions, K plays
a major role because plant membranes are highly
permeable to K and it is the most abundant cation in
plant tissues.
A knowledge of the functions of K in grape berries
will provide insight into how K requirements are deter-
mined and how excess K can be avoided. At present, little
is known about the precise functions of K in grape
berries. Nevertheless, the following information suggests
that K is essential for berry growth. The direction of K
transport is often towards the growing plant tissues, and
K is often redistributed from older to younger plant tis-
sues (Mengel and Kirkby 1987). The growth pattern of
grape berries (Figure 1) is described as a double sigmoidal
curve with an initial rapid increase in size (due to cell
division followed by cell expansion, in conjunction with
rapid accumulation of organic acids). This initial rapid
increase in size is followed by a lag period of slow or no
growth. The second phase of rapid growth is due entirely
to cell expansion and characterised by a rapid accumula-
tion of sugars (Coombe 1992). Inception of berry ripen-
ing, termed veraison, occurs during the transition from
the lag period to the second rapid growth phase. Veraison
is characterised by berry softening and colour changes,
and berries undergo major changes in chemical composi-
tion, including sugar accumulation and organic acid
depletion (Coombe 1992). A sharp increase in berry K is
also observed at the onset of ripening (Ollat and
Gaudillère 1996) (Figure 1).
Proposed explanations for the sudden onset of berry
expansion following veraison include ‘Sugar accumu-
156 Potassium nutrition in grapevines
lation’ (Coombe 1960), ‘Skin loosening’ (Considine and
Brown 1981, Huang and Huang 2001) and ‘Turgor-
driving’ (Zhang et al. 1997) hypotheses. Wall loosening is
the primary event initiating cell expansion (Cosgrove
1987). In grape berries, the increase in berry softening
(Coombe and Phillips 1982) and the decrease in skin
elasticity (Huang and Huang 2001) precede rapid berry
expansion at veraison. This suggests that the cell walls in
berry skin must loosen for berry expansion to begin.
Loosening of cell wall material involves the acidification
of the apoplast and the action of cell wall loosening
enzymes (Hager et al. 1971). Acidification occurs when
protons (H+) are pumped out of the cytoplasm and into
the apoplast by the membrane-bound ATPase (Hager et
al. 1971). Proton extrusion depends on the presence of K
in the apoplast since the uptake of potassium ions (K+)
balances the release of H+ to keep the plasma membrane
potential steady (Mengel and Kirkby 1987).
Both water and solutes (mainly sugars, K, malate, and
tartrate) contribute to the volume increase during cell
expansion. The water potential gradient is the driving
force for water movement (from high potential to low
potential). Decreasing cell turgor (pressure potential
becomes less positive) or increasing cell solutes (osmotic
potential becomes more negative) will result in the
decrease of cell water potential (becomes more negative)
because Ψw = Ψp + Ψs where: Ψw = cell water potential
(always negative); Ψp = cell turgor or pressure potential
(positive for living cells); and Ψs = osmotic potential
(always negative). Cell wall loosening lowers turgor,
which results in decreasing cell water potential and so
contributes to increasing the water potential gradient to
drive water entry. Increasing cell water content results in
increased turgor to drive wall extension but decreases the
water potential gradient. In order to maintain the process
of cell expansion, water potential gradient needs to be
maintained. A cell therefore may maintain the water
potential gradient by accumulating solutes. Sugar is the
major solute that accumulates in the grape berry during
the second rapid growth period, but K, as an abundant
and highly mobile element, can contribute significantly as
an osmotic component, especially in conditions of low
sugar accumulation (either by low production, transport,
Total soluble solids (°Brix)
or translocation). Other minerals contained in the berry
such as sodium, calcium, magnesium, copper, manganese
and phosphate also contribute as osmotic components.
However, the contribution of these minerals is small due
to their lower concentrations in the berry and because of
their lower mobility and/or their toxicity at high concen-
tration.
Translocation of solutes in grape berries is not clearly
understood (Ollat and Gaudillère 1996). The sink
strength of the berries for solutes during ripening could
possibly be controlled at the phloem unloading step
(Coombe 1992). Potassium may be involved in the
translocation of solutes into the berry through its roles in
phloem loading and unloading (Lang 1983). Because sug-
ars are the major soluble solids in grape berries, particu-
larly after veraison, total soluble solids (TSS), measured as
refractive index (oBrix) is often used as an indicator of
Australian Journal of Grape and Wine Research 9, 154–168, 2003
berry sugars. After veraison, there are large increases in
both K and TSS (Figure 1). A plot of K and TSS, mea-
sured in Carignane berries collected at 7 to 14 day inter-
vals between veraison and harvest, suggests that the rela-
tionship between K and TSS changes during berry matu-
ration and ripening (Freeman and Kliewer 1983). This
relationship has been described as the third degree poly-
nomial equation y = 11.44 + 5.87x – 0.41x2 + 0.012x3 (r2
= 0.96**) for irrigated vines (Freeman and Kliewer 1983).
When data of K and TSS for ripe berries from various
sources with variation in varieties and growing conditions
(Williams et al. 1987, Rühl et al. 1988, Bravdo and Naor
1996, Costantini et al. 1996, Walker et al. 1998, Esteban
et al. 1999, Cavallo et al. 2001) were plotted, variation in
the relationship between K and TSS increased and the
correlation coefficient of the relationship is low (r2 =
0.33) (Figure 2). A positive relationship between berry K
and berry sugar would be expected if K is involved in
solute translocation. Nevertheless, under conditions of
low sugar production, more K may be accumulated in the
berry possibly because K is a highly mobile osmoticum.
This may reduce the positive relationship between berry
K and berry sugar. Using TSS as the indicator of sugars to
investigate the relationship between berry K and berry
sugars may not always be accurate because other solutes
such as tartaric acid, malic acid and K itself also con-
tribute to the refractive index and therefore the TSS val-
ues. The relative contribution of sugars and non-sugar
solutes to the TSS values changes during different stages
along berry growth and may also vary depending on
growing conditions. Recent evidence, found in
Arabidopsis, maize and castor bean, suggests that a specif-
ic K+ channel may be linked to sugar unloading (Lacombe
et al. 2000, Bauer et al. 2000, Ache et al. 2001). More
studies to investigate the relationship between berry
sugar accumulation and berry K accumulation are
required.
r 2 = 0.33
K (mmol/L)
Figure 2. Relationship between K and total soluble solids of ripe
berries (r 2 = 0.33), using data from various studies with variation in
varieties and growing conditions (Williams et al. 1987, Ruhl et al.
1988, Bravdo and Naor 1996, Costantini et al. 1996, Walker et al.
1998, Esteban et al. 1999, Cavallo et al. 2001)
Mpelasoka, Schachtman,Treeby & Thomas
3. Root K+ uptake and translocation of K+ from
root to shoot
Potassium in the soil solution is taken up by plant roots.
Uptake of water and dissolved ions is restricted to root
tips. Movement of water and dissolved ions across root
cortex to endodermis occurs mainly through the apoplast.
Due to the presence of the casparian strip in the root
endodermis, water and dissolved ions have to move
across the plasma membrane into the root symplast (De
Boer, 1999). The two known systems of K+ transport
across membranes are ion channels (proteins which form
a selective pore in the lipid bilayer when open) and car-
riers or transporters (proteins that couple K+ transport to
the transport of a second ion with an energy gradient
more favourable than K+). It has been suggested that
external K+ status determines which mechanism is used
by a plant root, such that a sufficiently high external K+
concentration will allow uptake through channels, but
carrier-mediated uptake is necessary at a low external K+
concentration (Maathuis and Sanders 1996). However, it
has been demonstrated that K+ channels can also mediate
K+ uptake at µM external K+ concentrations when the
membrane potential is sufficiently negative (Hirsch et al.
1998). The identification and characterisation of K+ chan-
nels and K+ carriers (transporters) has been reviewed by
Czempinski et al. (1999), Rodriguez-Navarro (2000), and
Schachtman (2000); the regulation of K+ transporters by
Blatt (1999); and molecular mechanisms and regulation
of K+ transport by Véry and Sentenac (2003).
In the symplast, K+ diffuses from cell to cell through
plasmodesmata. To be loaded from the root symplast into
the transpiration stream, which flows through the xylem,
K+ has to cross the plasma membrane of cells within the
stele. The amount of K+ reaching the xylem is a highly
regulated process involving the activity of H+-ATPases,
channels and carriers. Centripetal diffusion can be driven
by a K+ gradient (i.e., cytosolic K+ activity of the stellar
cells is lower than that of cortical cells) and/or the differ-
ence in membrane potential of xylem parenchyma cells
and cortical cells (De Boer 1999). Translocation of K+ in
the xylem from root to shoot is regulated by the capacity
of the root in xylem loading and probably not by transpi-
ration (Tanner and Beevers 2001). Movement of K+ may
be altered by shoot demand because the shoot acts as a
sink for nutrients (Engels and Marschner 1992, Pitman
1972). Phloem also contributes to translocation of K+ and
is the major route of K+ translocation to growing tissues
such as developing leaves and fruits (Mengel and Kirkby
1987). Re-translocation from the shoot, via the phloem,
back to the root, where K+ is then reloaded back into the
xylem, may occur when the xylem K+ delivery to the
shoot exceeds shoot requirements (Drew and Saker
1984) and/or under K deficient conditions of the root
cells (Jeschke and Hartung 2000).
4. Berry K accumulation
Similar to many other growing tissues and storage organs,
grape berries are a strong sink for K. Potassium is by far
the major cation in ripe berries. For example, the con-
centrations (ppm) of inorganic cations in De Chaunac
Potassium nutrition in grapevines 157
berries at harvest were: K ≈ 2875, Na ≈ 200, Ca ≈ 100, Mg
≈ 110, Cu ≈ 2.2, and Mn ≈ 0.8 (Hrazdina et al. 1984).
4.1 Temporal pattern of berry K accumulation and berry K
partitioning
Berry K content generally increases over the season
(Conradie 1981, Possner and Kliewer 1985, Doneche and
Chardonnet 1992, Boselli et al. 1995, Rogiers et al. 2001)
with a sharp increase at the onset of ripening (Ollat and
Gaudillère 1996). In contrast, berry K concentration per
unit fresh weight may increase (Hale 1977), or remain
relatively constant (Boselli et al. 1995). This may occur
because berry K concentration is the result of berry
growth rate and the rate of K accumulation in the berry.
Berry K concentration will remain relatively constant if
berry growth and berry K accumulation are maintained
at similar rates. Berry K concentration will increase if rate
of K accumulated exceeds rate of berry growth. Factors
such as cultivars, crop load, climate and cultural practices
that affect rate of berry growth and/or rate of K accu-
mulation in the berry will affect berry K concentration.
These factors are discussed in section 5.
Information on the partitioning of K within a berry is
important for the development of better vinification
methods to reduce the negative influence of berry K con-
centration on wine quality and will serve as background
information for the study of the source, compartmen-
tation, and control mechanisms of the accumulation.
Among berry tissues, K concentration per unit fresh
weight is higher in the skin than in the fleshy pericarp
(pulp) (Coombe 1987, Iland and Coombe 1988, Walker et
al. 1998). The K concentration in the seeds is lower than
in the skin but slightly higher than in the flesh (Walker et
al. 1998). However, the degree of difference in K concen-
tration among berry tissues varies widely among varieties
and among rootstock/scion combinations. Results from
five grape varieties on own roots or on Ramsey (Vitis
champinii) rootstock show K concentration in the skin to
be 1.7–6.9 times higher than that in the pulp and 1.6–4.3
times higher than that in the seeds (Walker et al. 1998)
(Figure 3a).
The differences in K concentration among different
tissues may be attributed to differences in cell structure
and/or to the different roles K plays in different tissues.
Compared to pericarp cells, skin cells are smaller and
have thicker walls and more cytoplasm (Harris et al.
1968, Considine and Knox 1979, Nii and Coombe 1983).
The concentration of K in the cytoplasm is about 5 to 10
times higher than that in the vacuole (Flowers and
Läuchli 1983). While K concentration per unit weight is
highest in the skin, the weight distribution between
flesh, skin and seed of the berry is variable. Generally the
flesh weight is greater than the weight of skin or seeds
but this varies among different varieties and different
rootstock/scion combinations (Walker et al. 1998).
Therefore, the impact of skin K varies depending on skin
K concentration and the ratio of skin weight per berry.
This means percentage distribution of K between skin,
pulp and seed (Figure 3b), which integrates variation in K
concentration across different berry tissues and variation
158 Potassium nutrition in grapevines
K concentration (mg/g FW)
Percentage distribution of K
Figure 3. Potassium concentration of skin, pulp and seed (a) and
percentage distribution of K between skin, pulp and seed (b) for ripe
grape berries from own-rooted or Ramsey-grafted of Muscat Gordo
Blanco (M), Shiraz (S), Riesling (R), Cabernet Sauvignon (CS), and
Chardonnay (C). Values are means of three replicates with ten ripe
berries per replicate. For a, within each variety, asterisks represent
significant differences (P = 0.05) between own-rooted and Ramsey-
grafted. For b, within each berry tissue, values sharing at least a
letter (above the histogram) are not significantly different at P =
0.05. (Data taken from Walker et al. 1998)
in weight distribution of the tissues, is a better indicator
when comparing variety or treatment effects.
Within the pericarp, K concentrations are lower in the
peripheral zone and in the adjacent flesh than in the
more central tissue; this may be attributed to the pres-
ence of the functional vascular bundle and the nature of
phloem unloading (Coombe 1987). The longitudinal dis-
tribution pattern of K in a berry changes during the
berry’s development. When divided into four longitudinal
zones, K concentrations of green berries are higher in the
distal area near the stylar than those near the pedicel.
However, this pattern is reversed after veraison because
of a drastically reduced accumulation of K in the stylar
zone and a continuing accumulation in the zone close to
the pedicel (Possner and Kliewer 1985).
The only reported K+ transport mechanism in grape
berries is an ion channel. This channel is also expressed in
other tissues and is mainly expressed in berries before
Figure 4. This model shows type and location
of a K+ channel and two K+ transporters
localised in grape berry tissues. A SIRK K+
channel was found in whole berry and berry
pericarp (Pratelli et al. 2002). A KUP K+ • 2.21–3.29 mg K/g FW
transporter was found in seed and two KUP • One KUP K+ transporter
K+ transporters were found in berry skin
(Schachtman, Davies, Liu and Thomas
unpublished). SIRK encodes an inward
rectifying K+ channel in the Shaker family.
KUP encodes a specific type of K+ transporter.
Potassium concentrations in different berry
tissues are based on data from Walker et al.
1998. The berry image was redrawn from
Coombe (2001).
Australian Journal of Grape and Wine Research 9, 154–168, 2003
veraison (Pratelli et al 2002). We have identified two
a genes that encode potassium carriers that are expressed
in grape berries (Schachtman, Davies, Liu and Thomas,
unpublished) (Figure 4). Both are highly expressed
before veraison, with a low level of expression being
maintained after veraison. The expression of these carriers
is restricted to the skin, the tissue in the berry with the
highest K concentrations. Investigations on the other
types of K+ transport mechanisms in different berry tis-
sues and whether their activity changes during berry
b
growth would help to understanding the mechanism of
berry K accumulation.
4.2 Translocation of K into the berry
Since K is not metabolised, the amount of K accumulated
in a berry is expected to be equal to the amount of K
entering the berry minus that moving out of the berry.
Potassium movement occurs in both xylem and
phloem (Mengel 1976). In grape berries, the xylem
seems to be a minor route of K entry because xylem flow
into the berry may be low due to the low transpiration
rate of the berry. Rates of berry transpiration also
decrease during berry growth and development while
berry K increases. The decrease in berry transpiration is
probably due to decreased stomatal frequency (Blanke
and Leyhe 1987), degeneration of stomata to lenticels,
and the deposit of epicuticular wax (Blanke et al. 1999).
Apoplastic dyes, radioactive tracers, and girdling experi-
ments have been used to investigate translocation of
water and solutes in xylem and phloem. Results from
these studies, as outlined below, support the premise that
phloem is the major route of berry K entry.
Based on the assumption that K translocation occurs
in both xylem and phloem (Mengel 1976), whereas Ca,
which is phloem immobile, is translocated only in xylem
(Hanger 1979), changes in the K/Ca ratio in berries have
been used as an indicator of changes in the relative berry
K influx via xylem and phloem (Hrazdina et al. 1984,
Ollat and Gaudillère 1996, Rogiers et al. 2000, Rogiers et
al. 2001). While K accumulation occurs throughout all
growth stages, the accumulation of Ca ceases at veraison
in de Chaunac (Hrazdina et al. 1984) or continues at rates
much lower than K in Cabernet Sauvignon (Ollat and
Gaudillère 1996) and Shiraz (Rogiers et al. 2001). This
suggests an increase in the relative berry influx of phloem
K compared to xylem during post-veraison. Perfusion of
Skin
Seeds • 4.76– 8.82 mg K/g FW
• Two KUP K+ transporters
Whole berry & pericarp
• One K+ channel (SIRK)
Pericarp
• 1.29–2.88 mg K/g FW
Mpelasoka, Schachtman,Treeby & Thomas
apoplastic dyes (e.g. Eosin Yellow and azosulfamide)
(Findley et al. 1987, Creasy et al. 1993) and studies of
vascular anatomy (Düring et al. 1987, Findley et al.
1987) indicate the disruption of xylem in the berry after
veraison. Peripheral xylem elements in the brush zone
where the vascular bundles enter the berry are particu-
larly disrupted (Düring et al. 1987, Findley et al. 1987). It
has been suggested that the irregular stretching and
breaking of xylem tracheids in the peripheral bundles
observed after a rapid berry expansion post-veraison
might be the cause of this xylem disruption (Findlay et al.
1987). Dye studies using acid fuchsin, a synthetic xylem-
mobile and phloem-immobile solute, in Shiraz, Merlot,
and Muscat Gordo Blanco berries confirm the disruption
of xylem flow within the brush tissue, but the xylem in
the berry pedicel is still functional post-veraison (Rogiers
et al. 2001). More studies are required to confirm
whether xylem flow into the berry ceases totally after
veraison and whether this phenomenon varies among
grape varieties. This is because, after veraison, Ca ac-
cumulation in the berries does not always cease (Ollat
and Gaudillère 1996, Rogiers et al. 2000), and water
movement from vine to a berry with its pedicel girdled
(i.e. phloem interrupted) still occurs (Rogiers et al. 2001).
Blockage to dye movement may not indicate complete
cessation of xylem water flow. After veraison, water and
ions may still enter the berry through the xylem conduits
in the pedicel to the brush zone, then diffuse into the
pericarp via a non-vascular route (Rogiers et al. 2001).
A study on changes in berry weight and accumulation
of water and solutes of Shiraz berries also suggests
phloem sap as the major source of berry water and
solutes after verasion (McCarthy and Coombe 1999).
These authors also suggested that phloem flow into the
berry may be impeded after reaching maximum weight
resulting in pre-harvest weight loss and berry shrinkage.
Nevertheless, the continued accumulation of K and Ca
found subsequent to maximum berry fresh weight indi-
cates that a cessation of xylem and/or phloem flow to the
berry was not likely the cause for pre-harvest weight loss
during ripening in Shiraz (Rogiers et al. 2000). Whether
or not the cessation of xylem flow occurs during berry
ripening and/or the cessation of phloem flow occurs
after berries reach a maximum weight, there is clearly an
increase in the relative berry influx of phloem sap com-
pared to xylem sap during post-veraison. Through in-
direct evidence, Lang and Düring (1991) proposed that a
declining cell membrane integrity in the berry at the
onset of ripening which results in a breakdown of
apoplast:symplast compartmentation may also contribute
to this phenomena. The breakdown of apoplast:symplast
compartmentation decreases the water potential of the
phloem sieve tubes in the berry, increases the water
potential gradient of the phloem sieve tube between the
source (leaves and storage) and the sink (ripening berry)
and, therefore, stimulates phloem flow into the berry
(Lang and Düring 1991). The activity of invertase may be
involved because invertase transforms sucrose into two
hexoses. This transformation results in decreasing osmot-
ic potential in the berry and consequently increasing
Potassium nutrition in grapevines 159
water potential gradient that stimulates phloem flow
into the berry.
The rate of K entering a berry is likely to depend on
inflow volume of xylem and phloem sap and the concen-
trations of K in the sap. In grapevines, to our knowledge,
data on the flow velocity and sap concentration of xylem
is available for trunks, stems and leaves, but not for rachis
or pedicel levels. Potassium is the major cation in the leaf
xylem sap (Peuke 2000) and shoot xylem sap (Keller et
al. 2001). While leaf xylem sap K concentration did not
differ significantly with leaf age, leaf xylem sap flow is
highest in the middle leaves and lowest in the old and
young leaves. This may be attributed to differences in
assimilation and transpirational activities among different
leaf ages (Peuke 2000).
Quantification of the phloem sap flow is difficult due
to the extreme sensitivity of the phloem structure to
physical manipulation or wounding and the much lower
cross-sectional area (Köckenberger et al. 1997, Peuke et
al. 2001). Collection of phloem sap is also more difficult
than that of xylem sap and is subject to contamination.
The EDTA-facilitated exudation technique has been used
to collect phloem sap but quantitative assessment may
not be accurate. The use of aphid stylectomy to extract
phloem sap appears to be the most efficient technique as
it causes minimal mechanical damage to the plant and
provides a more accurate biochemical determination of
phloem sap components (Fisher and Frame 1984,
Girousse et al. 1991, Pritchard 1996). In grapevines, to
date, it has not been possible to use the aphid stylectomy
technique at the cluster level. Using the EDTA-facilitated
exudation technique, Glad et al. (1992) found a high K
concentration of phloem exuded from the peduncle of
Pinot Noir clusters (close to 800 nmol per exuded sample
collected in a 1.5 mL buffer solution during 4 h). In other
plant species K is the predominant cation in the phloem
sap (Ziegler 1975, Peuke et al. 2001) and is much more
abundant in the phloem sap than in the xylem sap (Pate
1975).
The inflow volume of xylem and phloem sap are a
function of flow velocity, cross-sectional area, and func-
tionality of xylem and phloem. Recalling that the amount
of K accumulated in a berry is a function of K concentra-
tion and inflow volume of xylem and phloem sap, unless
K concentrations in xylem and phloem sap are equal, rel-
ative contributions of xylem and phloem sap influxes,
which appear to change during berry development, will
affect berry K accumulation. This needs to be confirmed
through more experimentation. Exploration for method-
ologies to determine flow velocity and K concentration of
xylem and phloem at the cluster level will be very helpful
for such experimentation.
Loss of K from berries may occur through xylem
backflow from the berries to the vine (Lang and Thorpe
1989 and references therein) during the earlier stages of
berry growth. Reimport of this lost K back to the berries
may be possible if K is absorbed from the xylem apoplast
into the nearby phloem. It has been speculated that
xylem discontinuity after veraison might be a mechanism
to reduce the transport of apoplastic solutes out of the
160 Potassium nutrition in grapevines
berries (Findlay et al. 1987), which would otherwise
result from compartmentation breakdown in the berry
occurring by the onset of ripening (Lang and Düring,
1991).
4.3 Re-translocation of K from other parts of the vine into the
berries
The accumulation of K in permanent structures of the
vine (roots, trunk, and cordons) can take place through-
out the growing season, including the post-harvest period.
Potassium in these pools may be mobilised to support the
new roots, stems, leaves, and clusters when the uptake
from the soil is insufficient to meet the current demand.
During their development, berries appear to be the
strongest sink for K, especially between veraison and har-
vest. This may be due to the berry’s high demand for K
during rapid cell expansion. At harvest, clusters account
for 60% or more of the total K content of the above-
ground organs (Conradie 1981, Smart et al. 1985a,
Williams et al. 1987). Seasonal accumulation of K in dif-
ferent organs of a Chenin Blanc/99-R grapevine grown in
sand culture fed with Hoagland’s standard solution (con-
taining 4.7 mM K) shows that from veraison to harvest K
content in the clusters increases, and levels accumulated
are higher than the total amount taken up by the vine
(Conradie 1981) (Figure 5). During this period, K content
in the trunk, roots, shoots, and leaves decreases (Conradie
1981). This difference suggests that a significant amount
of K that accumulates in the berries during this period is
re-translocated from other organs. In this study, soil K
availability is not expected to be high because nutrient
solution holding capacity of the sand is low. Remobilis-
ation of K from other organs to the berries after veraison
is also reported in other studies (Downton 1977, Smart et
al. 1985b, Williams and Biscay 1991). In contrast to
these studies no K re-translocation was found during the
vine + bunches (notional data)
roots + trunk + shoots + leaves
roots + trunk + shoots
roots + trunk
roots
vine veraison
K content (g/vine)
flowering
bunches
budburst
Days following budburst
Australian Journal of Grape and Wine Research 9, 154–168, 2003
pre-harvest period in vines with a high K content (Lévy
et al. 1972). Remobilisation of K from other organs to the
berries may depend on soil K availability, K uptake
capacity of the roots, and rates of K translocation from
root to shoot to meet the berry demand for K.
5. Factors affecting berry K accumulation
Many factors may affect net accumulation of K by berries
through their effects on root K+ uptake, translocation of K
from root to shoot, re-translocation of K from shoot back
to root, the amount of K reserve, and the number of
berries and berry growth rates in relation to vine vigour.
These factors include soil, plant, vine microclimate and
cultural practice. Interrelationships among the effects of
these factors are likely to complicate any simple explana-
tion for the regulation of K accumulation in grape berries.
5.1 Soil factors
The uptake of K+ by plant roots is determined by the
levels of K in the soil that are available to plants, and by
the growth, distribution pattern and physiological activi-
ty of plant roots. These factors are influenced by physical
(texture, moisture, permeability, and depth) and chemical
(soil reaction, pH, and soil composition) properties of the
soil.
Not all of the K existing in the soil is available to
plants. Potassium in the soil exists in four forms, ranging
in the order of their availability to plants as solution >
exchangeable > fixed (non-exchangeable) > structural or
mineral (Sparks and Huang 1985, Sparks 1987). These
four forms of soil K are in dynamic equilibrium. The rate
and direction of the equilibrium reactions between these
forms, and hence K availability to plants, depends on the
rates of K+ uptake by plant roots and soil characteristics
such as mineralogy (types and amount of soil minerals
and size and degree of weathering of the mineral particles),
harvest
leaf fall
Figure 5. Seasonal
accumulation of K in
different organs of a
pruning grapevine (Chenin
leaves Blanc/99R) grown in sand
culture (Redrawn from
Conradie 1981)
shoots
trunk
roots
Mpelasoka, Schachtman,Treeby & Thomas Potassium nutrition in grapevines 161

moisture, pH and texture (Sparks and Huang 1985, Table 1. Potassium concentrations in expressed root sap, shoot
Northcote 1992, Horra et al. 2000, Zeng and Brown xylem sap and plant tissues (laminar, petiole, root) of ungrafted
2000). rootstocks, Dog Ridge and 140R, at different levels of K supply. The
unit of K concentration is meq/L for expressed root sap and shoot
Potassium ions, which are taken up by the cell rapid- xylem sap, and is % dry weight for plant tissues. Values are means
ly, generally compete strongly in cation uptake. However, of five replicates. Values across the row followed by different letters
when other cations are present in high concentrations, K+ are significantly different at P = 0.05 (Ruhl 2000).
uptake may be reduced. For example, K deficiency may
be induced in saline conditions where Na+ is the predom- Samples K concentration
inant cation (Chow 1990). At low external K+ levels, Dog Ridge 140R
uptake is highly specific for K+, while at higher external
K+ levels (> 0.5 mmol/L) Na+ can competitively inhibit K+ At 0.1 mM K supply
influx (Epstein et al. 1963). In addition, excess of Na+ and Expressed root sap 7.30 a 9.70 a
Cl- in saline soils creates high ionic imbalances that may Xylem sap 0.55 a 0.42 a
impair the selectivity of root membrane (Bohra and Apical laminar 0.93 a 0.74 a
Dörffling, 1993). Therefore, relative concentration of K+ Basal laminar 0.74 a 0.61 a
and other cations in the soil solution may be as important Petiole 1.38 a 0.82 a
for rates of plant K+ uptake as the concentration of K+ per Root 0.94 a 0.73 a
se. At 10 mM K supply
Expressed root sap 36.10 a 40.20 a
5.2 Variety, rootstock, and rootstock/scion combination Xylem sap 8.41 a 3.97 b
Plants differ in K efficiency, which is defined as their abil- Apical laminar 3.39 a 2.41 b
ity to obtain high relative yields at low K supply. For Basal laminar 3.16 a 1.90 b
example, cereals seem to be more K-efficient than dicots, Petiole 5.93 a 4.75 b
and sugar beet is more K-efficient than the potato Root 4.50 a 5.28 b
(Steingrobe and Claassen 2000). In grapevine, differences
in the K status of vines, berries, juice or must have been
reported when the following were compared: different George has higher capacity for xylem loading compared
varieties on own roots (Christensen 1984, Robinson and to 110R. When ungrafted rootstock varieties Dog Ridge
McCarthy 1985, Walker et al. 1998); the same varieties and 140R were compared at low (0.1 mM) K supply,
on own roots and on rootstocks (Rühl et al. 1988, Walker there were no differences in K concentration in expressed
et al. 1998, Gawel et al. 2000); different varieties grafted root sap and shoot xylem sap, or in root, petiole and
onto the same rootstock (Boselli et al. 1995, Arroyo et al. laminar tissues (Table 1) (Rühl 2000). At high (10 mM) K
1997, Walker et al. 1998); and different ungrafted root- supply, K concentration increased in all samples from
stocks (Rühl 1989b, 2000). both rootstocks. The K concentration in expressed root
Rootstocks have been used in many Australian vine- sap was similar for the two rootstocks but those in shoot
yards because of their nematode and phylloxera resis- xylem sap, petiole and laminar were higher in Dog Ridge,
tance and salt or lime tolerance (Rühl et al. 1988). Large while that in root tissue was higher in 140R (Table 1)
variations in the effects of different rootstocks on scion K (Rühl 2000). These data suggest that, at high K supply,
accumulation have been observed among rootstocks of 140R has a relatively higher ratio of root K accumulation:
different genetic origin as well as among those of the K translocation from root to shoot compared to Dog
same genetic origin (Rühl et al. 1988, Rühl 1991, Walker Ridge (Rühl 2000). There appear to be differences in
et al. 1998). V. rupestris rootstocks have a poor K+ uptake xylem loading and translocation from root to shoot
ability, while V. berlandieri rootstocks have a good to inter- among different rootstocks. Xylem loading of K+ is regu-
mediate K+ uptake ability, even at a low external K+ con- lated separately from root K+ uptake from the external
centration (Boselli et al. 1987). Berry K concentration solution (Engels and Marschner 1992). The activity of the
was higher for Sultana grafted onto Dog Ridge than that gene encoding K+ loading into the xylem can be regulat-
of Sultana grafted onto Ramsey, which are both selections ed by the stress hormone abscissic acid (ABA) (Gaymard
of V. champini (Hale 1977). These variations may be et al. 1998). More studies are required to determine
caused by differences in root K+ uptake capacity and/or whether variation in rootstock effects on scion K accu-
differences in xylem loading of K+ and translocation of K mulation is due to variation in the existence/expression
from root to shoot. of the genes encoding root K+ uptake or xylem loading of
Differences among rootstocks in rooting morphology K+ or due to variation in ABA production or the combi-
and density in the soil profile (Perry et al. 1983, Southey nation.
and Archer 1988, Swanepoel and Southey 1989) may Rootstocks also play an important role in the vigour
contribute to the differences in root K+ uptake capacity. and crop production of the scion. The effects of three
Comparing two rootstocks, Rupestris St George (or rootstocks, St George (V. rupestris), A×R #1 (or ARG1,
Rupestris du Lot, V. rupestris) and 110R (V. berlandieri × V. Aramon × V. rupestris Ganzin no.1) and 99-R (or Richter
rupestris), it was found that K+ uptake flux was similar, 99, V. berlandieri × V. rupestris), on vigour, crop production
but xylem sap K concentration was 40% higher for St and petiole nutrition of the scion were compared using
George (Swanton and Kliewer 1989), suggesting that St 22 scion varieties (Cook and Lider 1964). They reported
162 Potassium nutrition in grapevines
that St George produced the most vigorous scion growth
but the lowest fruitfulness per unit of growth, A×R #1
produced the highest fruit yields, but 99-R was the most
efficient in terms of fruit yield per unit growth. Average
for all scion varieties, petiole K concentration was similar
for A×R #1 and 99-R, but was highest for St George.
Increasing vine vigour or increasing crop production
may enhance K+ uptake and translocation as a result of
increasing demand for K. Increasing vine vigour also
increases the degree of shading in the canopy, which may
also enhance K+ uptake and translocation (see sections
5.4 canopy microclimate and 5.5.1 canopy management).
In terms of berry K accumulation, the same rootstock
may have variable effects on different scion varieties.
Berry K accumulation in five varieties, Muscat Gordo
Blanco, Shiraz, Riesling, Cabernet Sauvignon, and
Chardonnay, on own-roots or on Ramsey, was investi-
gated (Walker et al. 1998). Berry K concentrations were
higher in grafted vines than own-rooted vines for all vari-
eties except Chardonnay and was highest in Shiraz on
Ramsey (Walker et al. 1998). Rootstock/variety combina-
tions also influence berry weight (Hale 1977, Rühl et al.
1988, Walker et al. 1998) and skin/pulp ratio—and there-
fore berry K partitioning (Walker et al. 1998).
5.3 Berry growth, seed number, and phytohormones
It has been hypothesised that peach fruit growth is con-
trolled by hormones produced in the seed and that hor-
mones originating in the seeds or pericarp direct the
import of nutrients from other parts of the plant to the
fruit (Chalmers and van den Ende 1977). In grapevines,
increased seed number per berry results in a greater final
berry size (Cawthon and Morris, 1982, Boselli et al.
1995), but had little effect on berry K concentration
(Boselli et al. 1995). In two grape cultivars, Bacchus and
Madeleine, berry growth phases are influenced by num-
ber of seeds in the berry. The smaller the number of seeds
per berry the earlier the first growth phase ends and the
second and third growth phases begin (Staudt et al.
1986). However, the phenological relationship between
seed and pericarp development is highly variable among
seeded grape cultivars (Peynaud and Ribéreau-Gayon
1971). In one study, seed number had little relationship
with ABA or IAA contents in the berry (Cawthon and
Morris 1982). In another study, seed number per berry
was positively correlated with ABA and gibberellin con-
tent (Scienza et al. 1978). Although the roles of seeds on
nutrient accumulation are still unclear, it seems likely
that seed number per berry influences final berry size and
therefore berry K partitioning. Small differences in berry
size can lead to considerable differences in flesh:skin
ratios (Somers and Pocock 1986), with a higher percent-
age of skin per total berry weight in smaller berries. Skin
K concentration is also reported to be higher for smaller
compared to larger berries (Storey 1987) and this para-
meter may have an effect on wine quality.
The hormone ABA acts as a signal under water stress
to induce stomatal closing by opening outward rectified
K+ channels in guard cells, leading to K+ release, loss of
turgor and closing of stomata (Kearns and Assmann
Australian Journal of Grape and Wine Research 9, 154–168, 2003
1993). In grape berries a decrease in organic acids and an
increase in sugars are the major compositional changes
during berry ripening and have been found to coincide
with an increase in berry ABA at the onset of ripening
(Downton and Loveys 1978). If K+ channels are involved
in K accumulation in grape berries it is unlikely that they
would be regulated in the same way by ABA as are guard
cell K+ channels. This is because during ripening K is
accumulated (not released) by cells and this accumulation
coincides with ABA in berries reaching its maximum
(Downton and Loveys 1978) concentration.
ABA may however be important in regulating K
movement from roots to shoots particularly under stress-
ful conditions, which may impact the overall grapevine K
content and ultimately berry K content. In contrast to the
stimulatory effect of ABA on guard cell K+ release, in root
cells ABA reduces K+ efflux from stelar cells (Roberts and
Snowman 2000). This reduction in K+ efflux may lead to
less K+ being released into the xylem and less K trans-
location from roots to shoots. The underlying cause for
the reduced activity may be due to a reduction in expres-
sion of the gene coding for a particular type of channel
that is important in the release of K+ from stelar cells to
the xylem (Gaymard et al. 1998). From these studies it
can be concluded that ABA in roots may play a role in
regulating the long distance movement of K. If long dis-
tance movement can be shown to be important in berry
K accumulation, methodologies that can regulate root
ABA levels may be used as a management tool for con-
trolling berry K. One example would be mild water
stress such as partial rootzone drying (see also section
5.5.4), leading to increased ABA levels which may con-
tribute to lowering K movement from roots to shoots.
5.4 Canopy microclimate
Canopy microclimate, which is the climate within and
surrounding the canopy, plays an important role in deter-
mining grape and wine quality. The amount and spatial
distribution of leaf area and its interaction with above-
ground climate determine the canopy microclimate
(Smart et al. 1985b). Canopy microclimate is therefore a
result of the interrelationship between plant factors
(shoot density and vigour), climatic conditions, and
cultural practices (canopy management). The climate
parameters most influenced by canopy characteristics
are radiation, wind speed, and evaporative demand
(Smart et al. 1985b). The degree of light exposure, or
shading, and possibly evaporative demand appear to be
important in berry K accumulation. Increased accumula-
tion of K in vines, berries or juice has been observed as
the result of shading by artificial shade cloth (Rojas-Lara
and Morrison 1989) or by the canopy itself (Smart et al.
1985a, Archer and Strauss 1989, Dokoozlian and Kliewer
1995, Dokoozlian and Kliewer 1996). The effects appear
to be due to foliage shading and not cluster shading (Rojas-
Lara and Morrison 1989, Dokoozlian and Kliewer 1996).
The effects of shading on mean berry weight are
inconsistent: weight has been documented to decrease
(Rojas-Lara and Morrison 1989), increase (Crippen and
Morisson 1986), or was not affected (Haselgrove et al.
Mpelasoka, Schachtman,Treeby & Thomas
2000) due to shading. In contrast, berry sugar content has
decreased in almost all shading studies. Therefore we
speculate that the increased K translocated to berries may
be a mechanism to adjust osmotic potential for maintain-
ing turgor and water potential gradient when less sugar is
being imported to the berry in order to avoid or minimise
a reduction in berry growth.
5.5 Cultural practices
Several factors relating to soil, plant and vine microcli-
mate can be altered by cultural practices. For example,
irrigation and nutrient addition alter soil K mobility and
availability, and canopy management such as training,
trellising and pruning affect the vigour and microclimate
of the vine.
5.5.1 Canopy management
Canopy management is a technique that results in the
altered position and/or density of leaves, shoots, and fruit
to achieve a specific desired arrangement (Smart et
al.1990). The management of canopy density partly
determines the canopy microclimate. High shoot num-
bers and high vine vigour will result in high canopy den-
sity and an increased degree of shading within the
canopy. This can be regulated by three principal means of
canopy management: shoot number control, shoot vigour
control, and the use of a trellising/training system (Smart
1985). Canopy density, and hence degree of shading, can
be reduced by shoot pruning, by selective shoot removal
or by distributing the shoots over the wider areas.
On the basis of single-shoots, ‘high vigour’ implies
rapid shoot growth, with long internodes, thick shoot
diameter, large leaves, and more lateral shoots (Winkler
et al. 1974). Conditions known to reduce vine vigour
include water stress, high crop load, low nutrient avail-
ability, diseases and pests, root restriction, and the use of
low vigour rootstocks and/or scions. The effects of crop
load, nutrition and water stress on berry K accumulation
will be discussed in the following sections.
5.5.2 Regulation of crop load
Since berries are a stronger sink for K than other parts of
the vines, especially after veraison, it is possible that crop
load may affect the patterns of translocation and distrib-
ution of K within the vine. Crop load is usually defined as
the ratio of fruit weight to dormant pruning weight
(Bravdo et al. 1984) or of fruit weight to total leaf area
(Kliewer and Antcliff 1970, Kliewer and Weaver 1971).
Published data on the effects of crop load on berry K are
inconclusive. For example, in one instance cluster thin-
ning had no effect on juice K (Freeman and Kliewer
1983); in another, berry K increased when crop load
decreased (Hepner and Bravdo 1985). Crop load may
affect berry K accumulation by alteration of source/sink
balance for K. Crop load may also have indirect effects on
berry K accumulation through changes in the degree of
shading in the canopy, availability of assimilates and rates
of berry growth. Variation in the effects of crop load on
these parameters may depend on the proportion of and
the interaction between vegetative and reproductive
Potassium nutrition in grapevines 163
parts. Although, in principle, reduced crop load will
enhance berry growth, this effect will depend on the time
when the crop was thinned and the amount of crop
retained. Earlier thinning generally has a greater effect on
increasing berry growth and size compared to later thin-
ning. Overcropping may result in insufficient assimilate
supply, but a crop load that is too light may also reduce
assimilate availability for the berries due to increased
shoot growth, the competitive sink for assimilates.
Reduced photosynthetic rate under light crop loads was
also reported in apple as a result of carbohydrate accu-
mulation (Palmer et al. 1997, Wünsche et al. 2000).
There may also be a genotypic effect, i.e. different grape
varieties respond differently to crop load. Therefore, the
contradictory data published may be due to the use of
different grape varieties, different levels of crop load and
different time of thinning, different vine management, or
different weather and soil conditions during the study
period.
Excessive vegetative growth and increased shading is
generally associated with poor crop load (Bravdo and
Naor 1996). The ratio of fruit weight to effective leaf area
(i.e., the area of leaves exposed to light) and degree of
shading are more likely to be related to fruit composition
than the fruit weight to total leaf area ratio, which is the
common definition of crop load. The percentage of effec-
tive leaf area to total leaf area decreases when shading is
increased. Therefore, other factors, such as the training/
trellising system, may modify the effects of crop load
because they modify the degree of shading and so the
effective leaf area per unit fruit weight. Increased water
supply results in some similar effects to reducing crop
load such as increased shoot growth and increased fruit
weight (Esteban et al. 1999) (see section 5.5.4). Inter-
actions between irrigation and crop load treatments on
various parameters including fruit water relations and
growth and photosynthetic rate have also been reported
in apple (Mpelasoka et al. 2001). These factors need to be
accounted for when studying the crop load effects.
5.5.3 Nutrient addition
The effects of K addition on berry K accumulation are
variable. Results from some studies indicate that berry,
juice, or wine K increases when the plant is supplied with
K (Downton 1977, Morris et al. 1980, Rühl 1989a);
others did not find a significant effect of K fertilization
on berry or juice K (Rankine et al. 1971, Freeman and
Kliewer 1983). Variations in these findings may be attrib-
uted to the complicated chemical reactions of K in the
soil. Not all K that exists in the soil is available to plants
and it is unlikely that the increase in plant available soil K
is directly proportional to the amount of K fertiliser
added. The impact of K fertiliser on the level of plant
available soil K and on berry K accumulation is in-
fluenced by various factors, such as the amount and type
of fertiliser applied, the timing and frequency of applica-
tion, soil characteristics and management, the amount
and frequency of irrigation, plant root activity, and initial
vine nutrient status. Vines of different varieties, vigour,
and crop production may also respond differently to
164 Potassium nutrition in grapevines
specific levels of soil K availability.
Fertilisation programs are usually based on vine nutri-
tional status, which usually rely on the estimates of the
nutritional status of vegetative parts. In Australian vine-
yards, the petiole opposite the basal cluster at flowering is
commonly measured (Robinson et al. 1978, Robinson
and McCarthy 1985). Therefore, one important question
is the relationship between petiole K and berry K. In one
study, no relationship between petiole K and berry K was
found (McCarthy 1981), while others (Downton 1977,
Morris et al. 1980) found that berry K increased when
petiole K increased. The relationship between petiole K
and berry K is likely to change over the season due to
changes in sink strength and may also vary with different
crop levels, soil K availability, rate of uptake and trans-
location, and amount of K reserve in the permanent
organs. To avoid excess K concentration in grape berries,
berry K concentration should also be integrated into the
fertilisation program, and optimum K concentration
should be carefully calibrated in relation to growth, yield,
and juice and wine quality. A framework to estimate fer-
tiliser supply to achieve this aim is not yet available.
Besides K fertilisation, potassium metabisulphite is
sometimes used in the vineyard during the growing
season, at harvest and during vinification as a source of
sulphur dioxide to control diseases and microbial infec-
tion. However, the impact of potassium metabisulphite
application on the K levels of berry or wine has not yet
been quantified. Quantification of this impact is required
to determine if the use of other methods for controlling
diseases and microbial infection is necessary to achieve
desirable K levels and wine quality.
5.5.4 Irrigation
The effect of irrigation on berry K accumulation is not
widely documented. The available data suggest that
increased irrigation tends to increase berry K accumula-
tion. Berry, must and wine K concentrations were higher
for irrigated treatments than for non-irrigated treatments
(Freeman and Kliewer 1983, Hepner and Bravdo 1985)
in at least two studies. Increased K in the leaf blade
(Zaballa et al. 1997, Klein et al. 2000) and petiole (Klein
et al. 2000) under irrigation has also been reported,
which indicates a general increase in vine K status.
Potassium availability and uptake is reduced under con-
ditions of limited soil water (Dundon and Smart 1984).
Therefore, an increase in root uptake under irrigated con-
ditions may account for elevated vine and berry K accu-
mulation. Irrigation enhances the dissolution of K from
clay particles and its movement in the soil solution,
which facilitates supply to roots and higher uptake (Klein
et al. 2000). Generally, irrigation increases shoot growth
(Matthews et al. 1987, Williams and Matthews 1990,
Esteban et al. 1999), and therefore an increase in vine K
and berry K through irrigation may also be attributed to
increased canopy density and increased shading within
the canopy. Studies that integrate irrigation treatments
and pruning, training and/or shoot positioning treat-
ments will be useful to investigate the separate effects of
irrigation and canopy density on berry K accumulation.
Australian Journal of Grape and Wine Research 9, 154–168, 2003
Irrigation strategies that reduce the water supply to
grapevines may be used to reduce berry K accumulation.
However, a reduced water supply may induce water
stress to the plants, and severe water stress generally
reduces yield. The degree of stress depends on the
amount and frequency of the water supply, the evapora-
tive demand of the atmosphere, and the plants’ adapt-
ability to water deficit. Yield may be maintained or mini-
mally reduced with mild water stress, but the optimum
stress level is likely to vary among varieties and climate
conditions and is difficult to impose consistently and uni-
formly in the vineyard. Partial rootzone drying (PRD),
which was first proposed by Loveys (1991) for grapevines
as a possible control of vine vigour, creates the conditions
under which chemical signals can be provided by the dry
roots while the wet roots prevent the development of
plant water stress. There is evidence for positive effects of
PRD in improving canopy microclimate by reducing shoot
growth (Dry et al. 1995, Stoll 2000). Dry et al. (1995) also
reported a decrease in juice pH with PRD. Both outcomes
could relate to a reduction in berry K content due to PRD,
but to the best of our knowledge no published data on a
respsonse in K accumulation under PRD are available.
6. Conclusion and future research
During development, the grape berry is a strong sink for
K, especially after veraison. Excess K in grape berries can
have a negative impact on wine quality mainly because it
increases the pH of musts and reduces the colour quality
of red wines. Although pH can be adjusted during the
vinification processes, ensuring naturally low K levels
in the berry may be a more cost-effective method because
it involves reduced inputs and waste management.
Manipulation of berry K accumulation requires an
understanding of the function of K in the berries, how
berry K requirements are determined, and how K is
translocated into and out of the berry. Unfortunately,
information in these areas is still limited. Based on avail-
able information and the assumption that K may play a
role in berry expansion via osmotic and turgor regulation,
there are several potential methods for manipulating
berry K accumulation. These methods may include:
selective use of rootstocks to reduce root K uptake,
translocation of K from root to shoot, and scion vigour;
canopy management to alter the degree of shading and
the ratio of fruit weight to effective leaf area; and irriga-
tion strategies to reduce K availability to plants and to
manipulate plant root activity and vine water relations
and vigour. A genetic improvement program, either con-
ventional or using transgenic breeding, which targets the
production of K use efficient varieties (which have high
yields with a low K concentration in their dry matter),
may be beneficial. It is noteworthy that most, if not all, of
these suggested methods are interrelated in their effects.
Therefore, in addition to detailed effects of each method,
their interactions also need to be determined before
these methods can be recommended. Potassium is essen-
tial for vine growth and yield. Fertilisation programs
often aim for high yield and usually focus on K concen-
tration in the vegetative parts, using the petiole K level as
Mpelasoka, Schachtman,Treeby & Thomas Potassium nutrition in grapevines 165

Table 2. Examples of the areas related to grape berry K accumulation where useful information is lacking or unclear

Area Information lacking or unclear

Potassium and wine quality • Quantification of the effects of ‘time on skins’ and existing juice K concentration
on rates of additional K extraction during fermentation
Functions of K • Precise functions of K in grape berries and in different berry tissues
Berry K accumulation • Upper and lower limit of K required for different berry tissues
• Types and molecular mechanisms and regulation of K+ channels and K+ transporters
involved in berry K accumulation
• Quantification and control mechanisms of K concentration and flow velocity of xylem
and phloem sap at cluster level
Factors affecting berry K accumulation • Relationship between soil K, petiole K and berry K
• Control mechanisms for the influences of rootstocks on scion K accumulation
• Berry K accumulation in response to crop load, K nutrient addition
• Impact of potassium metabisulphite application (in the vineyard and in the winery) on K
levels of berry, juice and wine

an index. However, product quality appears to be a key
factor for market competition. Therefore, berry K con-
centration should also be considered in determining the
fertilisation program with optimum K concentration care-
fully calibrated in relation to growth, yield, and product
quality. Quantification for the effects of potassium
metabisulphite application in the vineyard and the win-
ery on the K levels of berry and wine is required to deter-
mine if the use of other methods for controlling diseases
and microbial infection is necessary to achieve desirable K
levels and wine quality.
Studies to quantify the effects of ‘time on skins’ on
rates of additional extraction of K from berry skins to
must and wine will help to identify the modification of
vinification methods for red wines to reduce the impact
of skin K. Examples of the areas related to berry K accu-
mulation where information is lacking or unclear are out-
lined in Table 2.
Acknowledgements
We thank Cathy Kromer and Janet Oriatti for their help
with the manuscript. We are grateful to Professor Paul
Kriedemann for his valuable advice on the manuscript.
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Manuscript received: 24 March 2003
Revised manuscript received: 7 August 2003