Amer J of Potato Res (2005) 82:301-307 301

Nutritional Requirements of Potatoes

D. T. Westennann

USDA-ARS, Northwest Irrigai ion and Soils Research Laboratory, Kimberly, ID 8:3341, USA
Tel: 208-423-6524; Fax: 208-423-6555; Email; dtwO`nwisrl.arsaisda,gov

ABSTRACT tigaciOn que se haga sobre nutriciOn adicional en plantas

geneticamente modificadas, agricultura de precision,
Plant nutrition is the practice of providing to the calidad alimentaria y seguridad, impurezas de los fertil-
plant the right nutrient, in the right amount, in the right izantes y otros aspectos de manejo deben ayudar signi-
place, at the right time. This paper gives an overview of ficativarnente al productor.
the roles that each of the 16 essential nutrients have in
plant nutrition, their relative mobility as related to defi-
ESSENTIAL NUTRIENTS
ciency symptom expression, and what is generally
known about nutrient responses to field applications on Only relatively few chemical elements are necessary for
potatoes (Solanum tuberosum L.) in the USA and plant growth. To be an essential chemical element from the
Canada. Maintaining high crop yields with minimum perspective of plant nutrition (a) it must be present for the
nutrient losses to the environment is and will continue plant to complete its life cycle, (b) its metabolic role cannot be
to be a significant challenge to the potato producer. replaced by another chemical element, and (c) it is directly
Additional nutritional research efforts in genetically involved in a metabolic process within the plant, either having
modified plants, precision agriculture, food quality and a direct role in the process or as a compound component
safety, fertilizer impurities, and other management con- involved in the process. The 16 chemical elements that fulfill
cerns should significantly help the producer in this these criteria are carbon (C), hydrogen (H), oxygen (0), nitro-
effort. gen (N), potassium (K), phosphorus (P), sulfur (S), calcium
(Ca), magnesium (Mg), zinc (Zn), manganese (Mn), iron (Fe),
RESUMEN copper (Cu), boron (B), molybdenum (Mo), and chloride (Cl).
The plant obtains three, C, H, and 0, from air and water, while
La nutriciOn vegetal consiste en proporcionar a la the remaining 1:3 are obtained from soil and fertilizer sources.
Manta el nutriente corrects, la cantidad correeta, el Nitrogen can also be obtained from the air by symbiotic organ-
lugar correct° y el momenta correcto. Este artIculo da isms for use by legumes and other plants.
una vision general de los roles que tiene cada lino de los It is only the intent of this paper to briefly descrihe the
16 nutrientes esenciales en la plants, su movilidad en role that each of the essential elements has in the plant, as they
relaciOn con la expresiOn de los sintomas de deficiencia are already fully described by others (e_g., Mengel and Kirkby
y lo que generalinente se conoce sobre las respuestas de 1979; Marscluier 1986). Carbon, hydrogen, and oxygen are
la aplicaciim en papa (Solarium. tuberosum L.) en el components of all organic compounds. Carbon is also a criti-
campo, en ELIA y Canada. El hecho de mantener cal component of the carboxylic group. Nitrogen is a primary
rendimientos altos con perdida minima de nutrientes en component of all nucleic acids, proteins, and amino acids.
el suet() es y eontinuara siendo un desafio signiflcativo Potassium is necessary for the activation of sonic enzyme sys-
para el productor de papa. Cualquier esfuerzo de inves- tems, the translocation of carbohydrates, and for osomoregu-
lation. Phosphorous is involved in the energy transfer process
Accepted for publication 12 January 2005. and is present in phosphorlated sugars, alcohols and lipids.
ADDITIONAL KEY WORDS: Sofano on htherosum, essential elements, Calcium functions as a structural component of cell walls, in
fertilization, tissue tests, research opportunities cell division and elongation, and membrane permeability. Mag-
302 AMERICAN JOURNAL OF POTATO RESEARCH Vol. 82

nesium is a component of the chlorophyll molecule and an enhanced uptake of one ion in response to the uptake of
essential cofactor for the phosphorylation process. Sulfur is a another ion, primarily to maintain electrical neutrality within
component of selected amino acids. Zinc is a cofactor for sev- the plant. Plants will partially compensate for this effect by the
eral enzyme systems, including dehydrogenases and involved production of organic ions internally or by releasing a H . or
in tryptophane synthesis. Manganese is involved in the photo- HCO3 ion into the solution surrounding the root_ The latter
synthetic process and as an activator for IAA oxidase. Iron is mechanism affects the availability of inorganic ions whose sol-
essential in electron transfer, for herne enzymes and chloro- ubility depends upon the pH in the rhizosphere.
phyll function. Copper is necessary for oxidase enzyme activ- Ions move to the root-solution interface by mass-flow and
ity and chloroplasts. Boron plays a role in cell wall stability, diffusion (Barber 1995). When ions are at a relatively high con-
cell differentiation and carbohydrate metabolism. Molybde- centration in the solution, there are sufficient amounts carried
num is essential for nitrate reductase and nitrogenase enzyme by the transpirational stream to supply plant needs, such as for
activity. Chloride functions in the photosystem II process and Cl, Ca, Mg. and NOrN. Mass-flow can also be important for
as an osmoticum. SO,-S and K. When ions are at a relatively low concentration
Other elements are classified as having a beneficial role in (<0.5 mg L-'), uptake is faster than movement by mass-flow
plants. These include sodium (Na), cobalt (Co), nickel (Ni), sil- and the concentration at the root-solution interface will be
ica (Si), vanadium (V), iodide (I), and selenium (Se). Sodium near zero. Under these conditions, movement to the soil-root
can partially substitute for K's metabolic role in some plants, interface is by diffusion down a concentration gradient. Diffu-
e.g., sugarbeets, and is considered to be an essential element sion is important for K, P, Zn, Mn, Cu, Fe, B, mid Mo. Some ions
for Halophytes. Cobalt is necessary for dinitrogen fixation and are taken up by direct contact of the root. with the ion on the
is a component of vitamin B 12 in legumes. Nickel was recently exchange complex or soil particle. The portion of ions gener-
shown to be a component of the crease enzyme system and ally taken up by this mechanism is not large since roots con-
necessary for ureide metabolism. Silica is necessary for the tact less than 5% the soil surface area.
growth of diatoms and stimulates the growth of some wetland Two additional factors affecting ion uptake are mycor-
grasses such as rice. Some algae species benefit from the pres- rhizae infection and the chemical, physical, and biological con-
ence of V. Plant species that act as Se accumulators some- ditions in the rhizosphere. Mycorrhizae are mutually beneficial
times show a growth benefit from Se additions. fungi that infect the root, extending the effective volume from
which plant roots take up nutrients. This process is most
PLANT UPTAKE AND MOBILITY important for ions that move to the root by diffusion. The
degree of root infection is also important. Today's commercial
Active and passive mechanisms are involved in moving potato varieties generally have a relatively low incidence of
ions from the solution contacting the roots into the plant cell. mycorrhizae infection. Infection also decreases as nutrient
The active process moves ions against an electrochemical or availabilities increase. As introduced in previous paragraphs,
concentration gradient and requires metabolic energy. The the immediate volume of soil surrounding the root, the rhizo-
passive process moves ions along an electrochemical or con- sphere, has an important role in nutrient uptake because nutri-
centration gradient and is generally considered to not require ent solubilities are dependent upon solution pH, which may he
metabolic energy. Most essential elements are taken up by a modified by root exudates. This area is also biologically active
combination of the two mechanisms, the active mechanism because of carbon enrichment from cell losses and root. exu-
being more important at lower solution concentrations. Sonic dates. The relative distribution of beneficial and harmful
ions are carried along with the transpirational stream. organisms in the rhizosphere and their effect on plant nutrition
Interactions can occur between ions during the uptake and health are largely unknown.
process. Competitive interactions occur between ions of simi- relative mobility of the essential elements in the
lar charge and size, e.g., K- and NH4* or NO3- and Cl-, by com- plant's vascular tissues affects the appearance of deficiency
peting for the uptake mechanism or carrier. The antagonism symptoms and nutrient application protocols. All nutrients are
interaction is similar to competition, but the ions can be dif- considered to be mobile in the xylem vessels. Xylem transport
ferent such as K. and Mr'. A synergism interaction is occurs in one direction, while phloem transport is bidirec-
2005 WESTERMANN: NUTRITIONAL REQUIREMENTS OF POTATOES 303

tonal. Since there is very little cross linkage, the mobility of plants generally take up more nutrients than required if avail-
the nutrient element in the phloem depends upon the form and able. Nutrient uptake is nearly complete when the majority of
the ability of the plant to load the element into the phloem. In tuber growth ends since little additional uptake occurs during
general, N, P, K, Mg, Cl, and S and their associated compounds the maturation growth stage (Westermann 1993).
are very mobile in the phloem, while Zn, Mo, and Cu mobilities Our relative ability to use a soil test to predict nutrient
are intermediate. Nutrient elements not mobile in the phloem requirement or a plant-tissue test to determine nutrient suffi-
of herbaceous plains are Ca, B, Fe, and Mn. The relative mobil- ciency or deficiency in the potato plant depends upon the
ity of Mn, B, and Cl in the phloem is partially dependent upon nutrient. In general, better information is available for plant-
the plant species. tissue tests than for soil tests for most nutrients (Table 1). The
Deficiency symptoms for phloem-mobile nutrients appear extremes of a nutrient deficiency can be easily determined, but.
initially on the older leaves, while deficiency symptoms for the the nutritional status of plants found in the transition zone
phloem-immobile nutrients appear on the immature leaves or between deficiency and adequacy is not always correctly
growing tips first. Complete correction of deficiencies for determined. There are wide ranges of known documented
phloem immobile nutrients is difficult with foliar sprays, par- field response data in the USA and Canada (Table 1). Known
ticularly for plants with the harvestable portion below ground. responses are well documented for N, P and K, while those for
Recent studies show significant B phloem mobility in plant S, Mg, Zn, and Mn are intermediate, and essentially none are
species that form B-sorbitol complexes (Brown and Hu 1996). available for Fe, Cl, and Mo. Only limited information is avail-
able for Ca, B, and Cu. Three states reported improved inter-
POTATO NUTRIENT nal tuber quality from applying Ca materials, but did not report
soil or plant calibration data for Ca. Limited data are also avail-
REQUIREMENTS
able for Mg and S. Of all the micmnutrients, reliable soil and
Potassium and nitrogen are found in the largest amounts plant data are available only for Zn, with only plant data for
in a potato plant, followed by Ca and Mg (Table 1). Most of the Mn. For the others, a significant amount of information is
phloem-mobile nutrients will be in the tubers at harvest while extrapolated from other geographic areas or other crops, or
the immobile nutrients will be in the residual vegetative por- sufficient nutrient concentrations are set by default because
tions of the plant. Total uptake amounts are site-specific since there were no responses to the applied nutrients. Tine. response
data reported in Table 1 are not inclusive as
TABLE 1—The relative whole plant nutrient uptake for a 56 Mg ha' tuber not all states responded to the information
yield, the general availability of a soil or plant diagnostic test request nor was the private consulting indus-
for each essential nutrient, and known field data available in try contacted. As management systems con-
the USA and Canada (uptakes in parentheses are estimates; tinue to improve, additional nutrient
states listed in parentheses have limited data on indicated deficiencies will be identified and reported.
nutrient). An example in another crop is a recent report
Total I iplake Diagnostic . Test Available Doeumentpd Responses & of wheat responding to Cl application in
Nutrient kg/ha Soil Plant. Calibration Data Available Montana (Engel et al. 1998).
N 235 yes yes USA & Canada Emerging nutrient diagnostic technolo-
P 31 yes yes USA & Canada gies include the chlorophyll meter and
K 336 yes yes USA & Canada
Ca 91 yes yes WI, (VA, WA, NY) remote sensing. The chlorophyll meter
Mg 63 yes yes CO, ME, NY, WI should be able to adequately assess the
S 22 yes yes CO, NE, WA, WI
Zn 0.12 yes yes ID, OR, WA, (ME)
plant's N status if it is properly calibrated.
Mn 1.00 no yes OR, NY, WI The user will have to recognize that many
Fe (2.0) 00 no
factors affect 11w plant's chlorophyll content
Cu 0.1 no no CO, WI
13 (0.2) no yes ME, WA when using the meter. Remote sensing may
Cl (2-3) no no eventually be a reliable diagnostic tool for
Mo (0.006) no no
the plant's real-time nutritional status, but it
304 AMERICAN JOURNAL OF POTATO RESEARCH Vol. 82

is doubtful if it will successfully be used to predict preplant roots are near the soil surface. This generally occurs when the
soil nutrient availabilities. The combination of remote sensing surface soil is always moist under the plant canopy. Other
and precision farming technologies has potential to increase application problems associated with fertigation are outlined
economic returns while protecting the environment. by Westenuann (1993).
Nutrients can be applied in various ways to meet. the
requirements for potato production (Table 2). Most nutrients FUTURE OPPORTUNITIES
can he successfully applied preplant if tilled into the rooting
zone before planting. Both Mn and Fe applied preplant may Agriculture is listed as a major non-point source con-
oxidize to unavailable forms before plant uptake, particularly tributing to the water-quality impairment problems of U.S.
on the high pH, calcareous soils. Nutrient source, e.g., chelated streams, rivers, and lakes (USEPA 1995). Runoff from agricul-
and inorganic salts, also influences the application method and tural lands contains dissolved organic and inorganic ions, and
rate for micronutrients. Application rates can generally be suspended solids that may contribute to water-quality prob-
lower when the chelate form is applied compared with the lems. Runoff nutrient concentrations generally increase as
inorganic salt. The nutrient should also be available for a their availabilities in the soil increase. Maintaining high crop
longer time interval after application when it is in the chelate yields with a minimum loss of nutrients to the environment is
form. a significant challenge. The following are selected opportuni-
The greatest benefit from a starter fertilizer material ties that could improve our ability to meet this challenge.
occurs when it is placed above the seedpiece because roots Genetic engineering has the potential to change the nutri-
develop at each node on the shoots above the seedpiece. Mate- tional relationships in the plant as known today. Potato plants
rials having a high salt. index should be avoided for use as with resistance to Colorado potato beetle and Roundup °-
starter fertilizers. Applications made post-plant are usually ready characteristics were being developed for public use in
done before row closure. When top-dressing, the fertilizer the late 1990s, but were then pulled from the market because
materials are broadcast on the surface, which could be fol- of public perception. We do not know if these changes altered
lowed by a final tillage operation, such as Inning. Side-dressed the plant's nutritional requirements. To fully realize their bene-
materials are usually physically injected with a shank into the fits and other changes, it may be necessary to know their
soil a few inches away from the seedpiece. Foliar sprays are effects on the nutritional requirements. These changes may
effective for most nutrients in correcting foliar deficiencies, have altered the plant's nutrient-uptake ability and/or the opti-
but not effective to correct tuber nutritional problems if the mum metabolic concentration within the plant tissue, which
nutrient is not mobile in the phloem. Fertigation can be an subsequently could affect the diagnostic soil and tissue-testing
alternative practice, particularly if the nutrient is mobile in the concentrations used for nutrient management. Additional
soil. A fertigation application of a soil-mobile nutrient (NOrN) genetic studies/modifications are also needed to improve the
can be more efficient than a preplant application when the disease resistance of the potato plant's root system and
nutrient is not leached out of the plant's root zone during the increase nutrient-use efficiencies. Nutrient-use efficiency
process (Westermann et al. 1988). When nutrients are fixed by would be significantly improved with more root hairs per unit
the soil, they should only be applied by fertigation when plant of root length, increased root growth longevity anti density,
and plants with greater rooting depth. This improvement alone
TABLE 2—Recommended fertilization practices for would significantly reduce the potential impact of potato pro-
potatoes. duction on water and environmental quality parameters, as
Fertilizer Application Nutrient well as reducing production costs. Nutrient-use efficiency
Preplant All might also he increased from improved nutrient utilization
Starter N, P, Zn, Mn, Ca, S within the plant via increased translocation or recycling.
Post-Plant
Development of plants with resistance to selected diseases
Top-dress N, P, 8, Ca
Side-dress N could also change their nutritional requirements, as there are
Foliar N, P, K, S, Ca, Zn, Mn, Cu, B, Fe close associations between disease resistance and nutritional
Fertigation N, P, K, C
adequacy (Huber and Graham 1998).
2005 WESTERMANN: NUTRITIONAL REQUIREMENTS OF POTATOES 305

Historically, soil fertility and plant nutrition researchers vitamins, carbohydrates, antioxidants, phytochemicals, and
have tried to eliminate all production variables except one digestibility of potato tubers. In addition, the potential for
when doing field studies. There are a few studies where com- tubers produced in soils used for disposal of animal manures
plex two-way interactions were thoroughly studied while or other by-products and biosolids to carry enteric organisms
there are almost no three-way interactions fully explored. The harmful to humans is not known.
single variable relationship can be expressed by the following large applications of fertilizers and soil amendments for
equation: potato production may cause the accumulation of heavy met-
als in tubers and eventually become toxic in the soil environ-
Y = f(x) (1) ment itself. Research activity has concentrated on cadmium
(Cd) since it is contained in many fertilizer materials (Anon.
where Y is the dependent variable (usually yield or nutrient 1998). In Australia, McLaughlin et al. (1997) found that fresh
uptake) in response to a single independent variable x (fertil- weight tuber Cd concentrations ranged from 0.004 to 0.232 mg
izer rate or soil test concentration or nutrient concentration in kg-', with a median of 0.033 mg kg-'. About 25.6% of the sam-
the plant) with a variance of ri,„1. All other variables were ples in their study exceeded the current ma yirman permitted
assumed to be constant. This process is used by the scientist concentration of 0.05 mg kg-'. An earlier U.S. market survey
to develop soil test correlation and calibration relationships showed a median Cd concentration of 0.028 mg kg -' for 297
used for recommending fertilization rates or to determine the tuber samples (Wu/Mk et al. 1983). The highest trace element
nutritional status of the plant. and heavy metal concentrations are found in sewage sludge,
Real-world production systems are much more complex rock phosphate, and phosphorus fertilizer samples compared
than that illustrated by equation (1). Within a given field there with other fertilizers or soil amendments (Raven and L,oeppert.
are both spatial and temporal variations in growing conditions. 1997). Canada and Washington State have already enacted a
This field variability is being partially addressed by site-spe- fertilizer law limiting the application of fertilizer materials on
cific or precision agriculture management practices. Ideally agricultural land based on their heavy metal concentrations.
under this protocol, plant nutrients for crop production would Similar laws are being considered in other states and nation-
be applied for the different production conditions within a ally (R. Stevens, WSU, pens comm). The actual solubility of
field. More than one production factor varies simultaneously heavy metals in soils and their assimilation by soil organisms
across a field and there is also the possibility that interactions and plants are urgently needed to adequately address these
occur between variables. The relationship expressed in equa- concerns since potato yield potentials may be limited in some
tion (1) then becomes production areas if fertilizer application rates are restricted by
law. In addition, their tuber concentrations and availabilities
Y = i((1) +./.(x:r) + +f(xi) +f(y,) + + vt. + (2) to the consumer must. be fully defined.
Nitrogen and phosphorus are the two major nutrients that.
Th ere is alnu ist no informal in on which to base dependable degrade water quality. Nitrogen as nitrate in drinking water is
nutrient recommendation rates under these production condi- potentially dangerous to newborn infants, causing methoe-
tions. The identification and quantification of the key variables rnoglobinemia, resulting in brain damage or even death. A limit
and their interactions will he necessary before the advantages of 10 mg L-' nitrate-nitrogen in water used for human con-
of precision agriculture will be fully achieved. This will not be sumption was set by EPA. Phosphorus contributes to the
an easy or inexpensive task. As well as being multidisciplinary, eutrophication of both freshwater and estuarine systems pri-
it will require the critical application of ninitivariable and other marily through increased algae growth. As such it is usually
advanced techniques (Mallarino et al. 1996). A creative exten- one of the targeted components for reduction in many total
sion of some of the concepts already available may be appro- maximum daily loads (TMI)Ls) for water-quality impaired
priate, e.g., DRIS (Sumner 1978) or crop-simulation models. streams on the 303(d) list..
There is increasing concern about the nutritive value of all Nitrate is highly mobile and can readily move below the
crops used for human consumption. Few field studies have crop rooting zone. Phosphorus is largely transported off-site
fully evaluated the effect nutrient elements have on protein, attached to the sediment, to be later released via dissolution or
;306 AMERICAN JOURNAL OF POTATO RESEARCH Vol. 82

made available when anoxic conditions are present. Nitrogen Researchers must become proactive to anticipate tomor-
must normally he added to achieve maximum economic row's needs as well as those of today. The ability to apply new
potato yields. Its efficiency may be substantially improved if it advances in technology from other fields as well as network-
is applied as close as possible to actual plant growth needs ing with others will be essential skills. These individuals will
(Westermann et al. 1988). Nitrate leaching may be reduced by also be required to do creative work with declining resources
improved irrigation management or a reduction in N fertiliza- in multidisciplinary environments to solve complex and diffi-
tion rates. The latter may also have the undesirable effect of cult problems since any new appropriate management prac-
reducing crop yields. tices must be sustainable and socially and environmentally
Phosphorus has more potential environmental impact acceptable. This will he a significant challenge for all who
when the available soil P concentrations are much higher than work in plant nutrition.
needed for plant growth. These concentrations are normally
found where manure or biosolids were applied based on the N ACKNOWLEDGMENTS
needs of the crop being produced. Phosphorus losses are
closely associated with soil erosion losses, but it can also The author wishes to thank all the individuals who pro-
move downward in the soil profile when the soil's sorption vided information on the documented responses and calibra-
capacity is saturated. There is also recent evidence that higher tion data available for potatoes within their states and Canada
P concentrations are found in the soil water moving in the by-
pass flow pores than in the bulk soil solution (Haygarth et al. LITERATURE CITED
1998).
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