Potato Research (2009) 52:305–317

DOI 10.1007/s11540-009-9145-2

Nitrogen Responses and Nitrogen Management

in Potato

J. Vos

Received: 20 March 2007 / Accepted: 24 December 2009 /

Published online: 27 January 2010
# EAPR 2010

Abstract Innumerable experiments have been carried out to establish the yield
response of potato to the rate of nitrogen (N) supply. Given the continuing change in
production level of potato and because of the need to maximise the nutrient use
efficiency and to reduce losses of harmful nitrogenous compounds to the
environment, such research is still necessary and topical. This minireview addresses
dose–response curves of fertiliser N input; the development of N fertiliser
recommendation systems; the so-called three-quadrant diagram of fertiliser N
response which dissects the ‘agronomic response’ into the underlying components;
the concept of critical nitrogen concentration as a function of crop biomass;
environmental aspects of fertiliser nitrogen supply; and the strategy of the potato
plant to cope with nitrogen limitation. European legislation sets limits on the input of
nitrogen and sets norms on water quality, making nitrogen use efficiency (NUE) a
critical issue. Precision agriculture may help to maximise NUE, provided an
adequate diagnostic system is developed that distinguishes between nitrogen
deficiency and other causes of spatially divergent crop performance.

Keywords Environment . Nitrate . Nitrogen . Nutrient use efficiency . Potato .

Precision agriculture . Solanum tuberosum . Yield response

Introduction

Ever since pioneers of plant nutrition like Carl Sprengel and Justus von Liebig (van
der Ploeg et al. 1999) discovered that plants absorb ions from the soil solution for
their growth, innumerable experiments were carried out to establish the yield
response of potato to the rate of nitrogen (N) supply. The reasons for this continued
interest in these dose–response relations was already implicitly discovered in the first

J. Vos (*)
Centre for Crop Systems Analysis, Wageningen University, Droevendaalsesteeg 1, 6708 PB
Wageningen, The Netherlands
e-mail: jan.vos@wur.nl
306 Potato Research (2009) 52:305–317

concepts that tried to articulate the nature of plant responses to the supply of
minerals. With his ‘law of the minimum’, Von Liebig (1840) already recognised
that a plant will positively respond to a larger supply of one particular mineral
factor as long as the one under question is the most limiting one. This means
implicitly that the response to N depends on the availability of other growth-
determining factors. The latter idea was expressed clearly in Liebscher’s ‘law of
the optimum’ (Liebscher 1895) stating that the production factor that is in
minimum supply contributes more to production the closer the other production
factors are to their optimum. Also, it became recognised that the response of the
potato crop to nitrogen supply is modulated by pests and diseases and by seed
quality, including its physiological age. The different durations of the crop cycle,
as expressed by maturity classes, is an obvious factor one might expect to affect
the N response too.
Since the time when von Liebig published his findings, the mean fresh potato
tuber yield increased, e.g., in the Netherlands from approximately 10 Mg ha−1 to
over 40 Mg ha−1 (Vos 1992). That increase reflects the development of crop
husbandry techniques including control of weeds, pests and diseases, improved
water management and of course adjustments in mineral nutrition, collectively and
interactively resulting in gradual increases in yield. Fertiliser recommendations
basically use recent historic experience on the response as best guess for the
response in the immediate future. In situations as experienced since the mid-1800s
where the average yield level gradually increased due to improved crop husbandry,
the recommendation is soon becoming obsolete and needs to be updated based on
new experiments.
It was not only the quantity of nitrogen supply that was addressed in research.
The effects were examined of the dominant forms of nitrogen, i.e. ammonium
versus nitrate, and merits were assessed of the chemical forms of compounds
(e.g. calcium nitrate, potassium nitrate, ammonium nitrate or ammonium
sulphate). The discovery that plants can absorb minerals via leaves led to
interest in foliar application.
The expansion of the processing industry (crisps, French fries) dramatically
changed forms of consumption of potato. Processing also set new criteria for quality
of tubers, including contents of total dry matter and of reducing sugars. Quality
demands from the industry triggered much research on the question of the role of
mineral nutrition, including nitrogen, as a determinant of quality parameters.
Over the last ca 30 years, environmental concerns have also kept the interest
going in new experiments addressing the N response. A new perspective became the
fate of nitrogen not absorbed by the crop. The possible fates include:
(a) immobilisation in the soil;
(b) gaseous emission to the environment via volatilisation as NH3 or via
denitrification yielding N2 and traces of N2O;
(c) washing out as NO3− via leaching; or
(d) carryover to the next cropping season as mineral N in the soil.
The only emission route without environmental consequences is gaseous loss of
N2. National and international legislation and targets on environmental quality
dominate current research on supply of N to crops. In Europe, very influential
Potato Research (2009) 52:305–317 307

documents are the Nitrate Directive (1991) (91/767/EEC) and the Water Framework
Directive (2000) (2000/60/EC). EU member states need to implement national
legislation so as to comply with the environmental aims specified in the directives.
As nitrogen input is basically limited to a maximum level, this shifts the focus of
research from finding the optimum rate of input to how to best make use of the
permitted maximum amount of external supply of N.
Organic manures have regained an important position in crop nutrition. The use
of such sources and the environmental issues have triggered many questions on rates
of transformation and transport processes involving N in the soil–plant–atmosphere
system. These processes include mineralisation (ammonification) of N from soil
organic matter and from incorporated crop residues, manures and amendments;
immobilisation of mineral N in microorganisms when the C/N ratio of substrate for
growth is too low for microorganisms to synthesise their body compounds;
nitrification, i.e. the microbial conversion of NH4+ to NO3−; denitrification, i.e. the
microbial conversion of nitrate into N2 and traces of N2O under anaerobic
conditions; atmospheric deposition of N compounds, primarily related to traffic
and ammonia emission from animal production sites; and crop uptake as related to
root growth and functioning and soil environment. Basically, the domain of research
extends to the whole N cycle looking for ways to reduce environmental pollution
and maximising plant uptake and plant production from a limited amount of external
N input. Disciplines involved include soil biology, soil physics, plant physiology,
agronomy, atmospheric chemistry and meteorology.
Recommendation systems for N nutrition were developed relatively recently in
comparison to recommendations for other macronutrients such as P and K. This
is explained from the many processes affecting the availability of N, inherently
making the estimate of an optimum (future) rate a tricky business. Initial N
nutrition recommendation systems delivered one single recommended rate of
supply at the start of the growing season. Later, it was recognised that such
recommendations stand high risks of being wrong because the required external
input depends on crop growth (i.e. the actual demand for N) and on supply from
sources other than the external input. Both unknowns at the beginning of the
season are affected strongly by the weather in the season to come. One way of
improving recommendations is to reduce the horizon of prediction from a whole
growing season to part of the growing season, i.e. the introduction of split supply
systems (Vos 1999). The basic philosophy is to start with a rate of supply that is
probably limiting. Subsequently, on one or two occasions during the growing
season, an assessment is made of the N status of the crop, and this N status is
converted into a recommended action for additional, in-season N supply.
Disciplines involved here include physiology, remote sensing and physics, as the
more promising systems derive an index for N nutrition status from analysis of the
reflection spectrum.

Nitrogen Response Curves

Von Liebig postulated a linear yield response to a larger rate of supply until a factor
other than N would become limiting. In 1909, Mitscherlich was probably the first to
308 Potato Research (2009) 52:305–317

express dose–response curves in mathematical formulae (Mitscherlich 1909). He

proposed a negative exponential equation to describe the diminishing return. Written
as a derivative, this reads:
dY =dNa ¼ aðYmax
Y Þ ð1Þ
which means that the degree of increase in production with an increase in the
amount of N applied, Na, is at any point proportional to the amount by which the
actual yield, Y, falls below the upper yield limit, Ymax; α is the proportionality
factor.
The integrated form is:


Y ¼ Ymax 1
e
aNa : ð2Þ
Mitscherlich’s law captures the general economic experience of diminishing returns
in response to ever larger levels of input. Yield asymptotically approaches Ymax,
implying denial of the existence of an optimum rate of N supply above which a
decrease in yield occurs. Indeed, a strong decline in fresh potato yield with an
increase in N input is rarely found (Fig. 1).
Yet, in view of practical recommendations, it is important to discern an economic
optimum taking the price of the produce and the cost of fertiliser application into
account. In general terms, the optimum is the point where the value of the first
derivative of the response function (no matter its mathematical form) equals the price
of application of one unit fertiliser. Neeteson and Wadman (1987) reevaluated results
from 99 fertiliser trials and described with two different models. Their first model
was a second-order polynomial:

Y ¼ b0 þ b1 N þ b2 N 2 kg ha
1 or Mg ha
1 : ð3Þ

70
fresh tuber yield (Mg ha-1)

60

50

40

30
0 100 200 300 400
N application (kg ha-1)

Fig. 1 Fit of the Mitscherlich model (Eq. 2) fitted to experimental data on fresh tuber yield (open squares)
in response to fertilizer N application rate (full drawn line); the equation for the fitted line is Y=38.4 + 29
(1 − e−0.0053X) R2 =0.984. Note that an intercept (38.4) was added since the original Mitscherlich model
passes through the origin. The hardly visible broken line represents a fit using a second-order polynomial
(Eq. 6) with equation: Y=38.8 + 0.125X − 0.00016X2, R2 =0.983. Data from 1988 in Vos (1997)
Potato Research (2009) 52:305–317 309

The point of optimum N input, Nop, was the level where

b1 þ 2b2 N ¼ P ð4Þ
or rewritten explicitly:


Nop ¼ ðP
b1 Þ=2b2 kg ha
1 ð5Þ
where bx are coefficients and N is the rate of N application and P the cost of
application per unit N.
Their second model was a modified exponential equation:


Y ¼ b0 þ b1 N þ b2 eaN kg ha
1 or Mg ha
1 ð6Þ
with the optimum rate of supply given by:
Nop ¼ ðlnððP
b1 Þ=ab2 ÞÞ=a: ð7Þ
Neeteson and Wadman (1987) concluded that the modified exponential equation
fitted the data best and tended towards lower values for Nop than the second-order
polynomial. However, it is a weakness of their dataset that data points are missing
between 0 and 100 kg ha−1, the latter being near the optimum in several cases.

Fertiliser Recommendation Systems

Fertiliser response trials provide an indication of the ‘average’ N requirement of the

crop. Since the initial amount of mineral N varies in the soil profile at planting, the
first recommendation systems accounted for mineral N measured in spring (Geijpens
and Vandendriessche 1996). The depth of sampling is an issue here; for starch potato
in the Netherlands, for instance, sampling depth was limited on statistical grounds to
the top 30 cm, whereas 0–60 cm proved better for ware potato on clay soils, yielding
the rule:


Na ¼ 270
1:1 Nmin kg ha
1 ð8Þ
where Nmin is the amount of mineral N (NO3− and NH4+) extracted from the 0- to
60-cm layer. There was enormous scatter in the data yielding this recommendation
rule, yet this was the best there was.
Of course it was recognised that recommendation systems such as captured in
Eq. 8 take only one factor into account, i.e. initial soil mineral N. More elaborate
systems define the N requirement, Nreq, as the difference, or the ‘balance’ between
sources of N, Nso, and sinks for N, Nsi (i.e. processes removing N from the pool of
available mineral N in the soil), where the estimated crop N uptake is included in the
sinks (Rémy 1981; Goffart et al. 2005; Wu et al. 2007):
Nreq ¼ Nsi
Nos : ð9Þ
The sources include initial soil mineral N and mineralisation of organic matter
during the growing season. The latter is dependent on the organic matter content of
the soil and the rate of addition of fresh organic materials (crop residues and manure)
and the C/N ratio of added organic material. Dry and wet atmospheric deposition can
310 Potato Research (2009) 52:305–317

assume an importance that cannot be ignored. Tables have been compiled based on
experience and calculation that specify subtractions from the standard recommended
rate to allow for in-season mineralisation. Other sinks are the estimated leaching
losses between soil sampling and the time of fertiliser N application.
For long, it has been standard practice to apply one single dose of nitrogen
fertiliser, either in the seed bed, or shortly afterwards. However, in view of the
difficulty to assess at planting the magnitude of the future seasonal demand and the
future seasonal supply from sources other than the fertiliser, split application has
aroused interest. In this way, the time horizon of the decision is reduced and one or
more options are created to correct the situation. The idea is to supply, e.g. 60–70%
of the anticipated requirement in the seed bed and decide on the application of the
remainder in the period from approximately mid-June to mid-July (depending on
maturity class). In the ‘Kulturbegleitenden Nmin Sollwerte System’ (Lorenz et al.
1985) and in similar systems in other countries, the soil mineral N is sampled during
the season and provides the basis for decision on supplementary nitrogen. That
calculation can pertain to the period from the point of decision to harvest or to a next
anticipated soil sampling date prior to harvest. Standard curves for the uptake of N in
the crop over time (e.g., Breimer 1989) provide an estimate of the anticipated N
uptake in a period of time between two soil samplings. The basic idea is that N
requirement of a crop is relatively conservative over years (although it can be
adjusted to the conditions in a certain year, e.g., after an episode of drought), whilst
all other transformation and transport processes would reflect in the level of soil
mineral N. If uptake in a particular part of the season is less than ‘average’ or
mineralisation is very rapid, these would be mirrored in high soil mineral N that can
be accounted for when calculating the N supply for the next period. The system was
first developed for field-grown vegetables but has not gained much application in
potato production.
There has been intensive search for other methods to assess the nitrogen status of
crops in the season. These include nitrate concentrations in sap extracted from
petioles or stem ends (Nitsch and Varis 1991; Gianquinto et al. 2004/2005), with do-
it-yourself kits consisting of test strips and a colour density metre. Chlorophyll
measurement with handheld equipment (e.g., SPAD 502 of Minolta) is also regarded
as a relatively quick and easy method to assess the N status of potato foliage
(Jongschaap and Booij 2004).
The reflection spectrum of the crop contains information on properties of the crop
(Jongschaap 2006) that can be converted into indices such as crop biomass, leaf area
index or crop N content.
Methods based on soil mineral N, sap nitrate concentration or chlorophyll index
or reflection measurements all have in common that a test result needs to be
interpreted and converted into a recommendation for action. In principle, one needs
to determine the effects of several doses of additional nitrogen for a broad range of
test values and extract from such data an interpretation of the test result.
Computer models are expected to be able to play a role in supporting crop
management decisions. One may expect reasonable guidance from computer models
that simulate the course of events in the soil and in the crop. It is especially
important that at some points in time, the simulated course of events can be forced to
agree with the one as derived from observation such as reflectance, e.g., measured
Potato Research (2009) 52:305–317 311

with a Crop Scan (Jongschaap and Booij 2004; Jongschaap 2006). Although
scientifically a sound approach, model-guided decision support has not gained wide
support yet, perhaps because too little effort is made to calibrate the model
(particularly the soil module describing N transformations and N transport) for the
local conditions (van Alphen 2002). Daisy (Abrahamsen and Hansen 2000) and
WAVE (Ducheyne et al. 2001) are examples of models that simulate nitrogen
transformation and transport processes in the soil–plant system. However, as yet
models do not feature as components of (commercial) systems generating
recommendations for interventions in the season.

The Three-Quadrant Diagram

The response of tuber yield to the rate of N supply is called ‘agronomic response’.
Even on a farm or experimental plot with constant management, the agronomic
response curve differs from year to year. That yearly variation indicates the room
that exists for better targeting the N application using smart decision support
systems. The three-quadrant diagram (van Keulen 1982; Vos 1997) provides a useful
tool to analyse the background of year/site/cultivar-related variation in agronomic
responses (Fig. 2). The agronomic response (quadrant I) is split up into a ‘soil and
root-related domain’, i.e. quadrant III, and a physiological response, i.e. quadrant II,
showing the amount of tubers that the plant manages to produce for a given amount
of N taken up from the soil. The factors that are primarily responsible for the value
of the intercept of the relation in quadrant III are the amount of soil mineral N at
planting (before fertiliser N is added), the rate of N mineralisation during the season
and contributions of N from other sources such as atmospheric deposition and
biological fixation. Losses during the growing season, e.g., due to leaching and

Fig. 2 Example of a ‘three- 70 tuber yield (Mg ha-1)

quadrant diagram’ of response
to fertiliser N application in
potato. Quadrant I is the ‘agro-
nomic response’, i.e. tuber yield
in response to rate of N applica-
tion. Quadrant II is the ‘physio- quadrant I quadrant II
logical response’, i.e. the tuber
production per unit of N taken up
in the crop. Quadrant III shows the N application (kg ha-1) N uptake (kg ha-1)
relation between total N uptake
(tubers plus haulm) at maturity 300
400
and the rate of N application. The
properties of the curve in quadrant
III is determined by processes in
the soil and root growth and root
quadrant III
functioning. The agronomic re-
sponse follows from the responses
in quadrants II and III. Data from
Vos (1997): solid squares 1988,
open squares 1990 N application (kg ha-1)

400
312 Potato Research (2009) 52:305–317

denitrification, equally affect fertiliser nitrate and nitrate from other sources and will
affect both slope and intercept. Ammonia volatilisation at the surface only affects the
slope. Insufficient root proliferation is likely to reduce the intercept and reduce the
slope (smaller fraction uptake per unit applied).
The slope of the line in quadrant III represents the apparent nitrogen recovery
(ANR) defined by:


ANR ¼ Nf
N0 =Ns ð10Þ
where Nf is the N uptake in a fertilised plot, N0 the uptake in control without
fertiliser and Ns the rate of supply of fertiliser N.
Neeteson et al. (1987) showed a continuous decline of ANR with an increase in
rate of supply. Greenwood and Draycott (1988), working with other vegetables than
potato, argued that the upper value of ANR for arable crops and vegetables is about
0.7. In cereals and grasses, ANR of 0.7 is maintained until the point where supply
exceeds the uptake capacity of the crop, beyond which point ANR is bound to
decline for higher rates of supply. Vegetables, representing potato as well, in
contrast, do not show constant ANR initially, but continuous decline of ANR for
each additional unit of N input. Greenwood and Draycott (1988) attributed this
primarily to the differences in root densities early in the season between cereals and
grasses on the one hand and vegetables and potato on the other hand. Some of the
fertiliser N applied to potato will be remote from the roots for considerable time
before being absorbed by the roots. This would inevitably lead to larger losses from
soil than in cereals and grasses.
Because of the monotonous decline in ANR, Vos (1997) described the curves in
quadrant III with a second-order polynomial and suggested that the coefficient of the
linear term represents the initial ANR and the value of the quadratic term the rate of
decline in ANR for each additional unit of N input. He also implied an inverse
relation between the intercept and the linear coefficient, but this was not
substantiated with data.
The amount of useful plant product produced per unit of N present in the whole
crop is commonly called the nitrogen use efficiency (NUE). The factors responsible
for the position and curvature of the line in quadrant II include the harvest index for
dry matter and the contribution of each unit of N resident in vegetative parts to crop
photosynthesis. Genetic variation is supposed to exist and is being searched for.

Greenwood’s Concept of Critical Nitrogen

Intuitively, one can surmise that a relation is likely between the current nitrogen
concentration in the crop and its rate of growth. Greenwood and co-workers
addressed this question extensively and came up with the concept of critical N
concentration, Ncrit (Greenwood et al. 1990; Gastal and Lemaire 2002):

Ncrit ¼ 5:7 W
0:5 ð%Þ ð11Þ

where W is the crop dry aboveground biomass expressed in megagrams per hectare.
As an approximation, Eq. 11 is valid for all C3 crop species, although some gain in
Potato Research (2009) 52:305–317 313

precision arises from crop-specific determination of Ncrit. Ncrit represents the

minimum N concentration in plant material of a given total biomass that is needed
for unrestricted growth. Ncrit declines for larger W, i.e. as the growing season
advances, because the fraction of tissue with low N concentrations assumes a larger
proportion of the total biomass in older than in younger crops. By definition, the
growth rate drops below the potential if for a particular magnitude of the biomass, W,
the actual N concentration, Nact, is lower than Ncrit. Colnenne et al. (2002) defined
the nitrogen nutrition index (NNI, no dimension), by:
NNI ¼ Nact =Ncrit : ð12Þ
Colnenne et al. (2002) showed in oil seed rape that seasonal, weighted NNI
correlated well with the relative yield, the latter being maximal if the seasonal
NNI was close to unity. Such data indeed reinforce the validity of the concept
of a critical nitrogen concentration that needs to be achieved for unrestricted
growth. The determination of Ncrit, involving sampling the biomass and
laboratory analysis of the nitrogen concentration in the dry matter, could provide
a basis for decision support on in-season additional N application. However, the
required sampling and analyses make the method impractical. That stimulated the
search for indirect methods that are quick and cheap to apply on large areas of
potato.
Equation 11 was found to be approximately valid for potato too. The statement
that the coefficients of Eq. 3 are largely similar across C3 species of very different
morphology implies that genetic differences in NUE among genotypes within a
species are not likely to be large.

The Nitrogen Response of Growth and Development

The growth period of the crop can be divided into three phases (Fig. 3), i.e. the
period from planting or emergence to full soil cover and maximum light interception
(phase I), the period of maximum light interception (phase II) and the period of
decline in light interception and soil cover, reaching zero at maturity (phase III).

Fig. 3 Fraction radiation inter-

ception, practically equivalent to
fraction of soil covered with green
plant material, as a function of
time after planting for three rates
of fertiliser nitrogen supply: zero
N control (circle), 50 kg ha−1 N
(triangle), 200 kg ha−1 N
(diamond) and 400 kg ha−1 N
(square). Data from 1990
in Vos (1997)
314 Potato Research (2009) 52:305–317

Plant yield increases with a larger rate of supply of N until some maximum yield is
reached. In general, more yield in response to more N can result from:
(a) a larger crop growth rate over the same total crop duration;
(b) similar average crop growth rate over an extended period of growth; or
(c) from a combination of (a) and (b).
Higher crop growth rate can arise from shorter durations of phases I and III and
longer duration of phase II. In potato, nitrogen has comparatively little effect on
phase I. The duration of phase II and whether or not full soil cover is attained is very
much affected by N supply. Apparently, N affects the rate of senescence only
marginally. The total growth period is prolonged for larger rate of N supply because
phase II extends with better N nutrition.
Such responses to low or high supply of N arise from effects of N on rate and
duration of appearance of leaves and branches on the potato plant and depend on the
active life span of individual leaves. Many processes determining leaf production are
not sensitive to N over a wide span of (suboptimal) N supply. These include the rate
of leaf appearance, the timing of appearance of basal or apical lateral branches and
the duration of leaf expansion. Specific leaf area is not systematically altered by N
over a large range of supply. For spaced plants, the active life span of leaves tends to
be longer at high rates of N supply than at lower rates (Biemond and Vos 1992).
Full-grown areas of individual leaf ranks are extremely sensitive to N supply
because leaf expansion rate is sensitive, but the duration of leaf expansion is not. In
conclusion, the prime factors responsible for divergent patterns (Fig. 3) are the
number of branches per plant and individual leaf size Vos and Biemond (1992).
For a large range of N supply, the leaf/stem ratio (dry weight) remains constant
and so do the harvest indices for dry matter and nitrogen of fully matured crops.
However, when high-N crops are prematurely harvested and compared to low-N
crops, the harvest indices are higher for the latter than for the former. There is some
delay in the onset tuber bulking with large rates of N supply. At high rates of N
supply, there is some shift in relative N distribution in the plant in favour of the
proportion N in stem material (Wu et al. 2007).
The nitrogen concentration in leaves and its change over the life span of the leaf area
are not very sensitive to N nutrition. Since photosynthetic capacity is associated with
leaf N concentration, it follows that photosynthetic capacity of leaves is only weakly
affected by N nutrition (Marshall and Vos 1991; Vos and van der Putten 1998, 2001).
These observations led to the proposition that potato adapts its foliar development to
limiting N supply in such a manner as to maintain productivity per unit leaf area whilst
adjusting total leaf area per plant (through leaf size and branching). In other words,
light interception per plant varies in response to N, but intrinsic productivity per unit
leaf area does not. Maize, for instance, shows the opposite strategy: under N
limitation, the plant puts priority on maintaining leaf area (i.e. maintaining light
interception per plant) to the detriment of leaf N concentration and associated
photosynthetic capacity (Vos et al. 2005). Potato does not ‘dilute’ leaf nitrogen in
N-limited conditions, whereas maize does so. It might be postulated that the radiation
use efficiency is insensitive to N supply in potato and sensitive in maize.
The ‘potato strategy of response to (limiting) N supply’ implies that plant biomass
is responsive to N whilst N concentration in leaves is conservative. Total nitrogen
Potato Research (2009) 52:305–317 315

and nitrate-N of stems and petioles are more responsive to N than leaf N. This means
that methods to monitor the N status of the crop that directly assess the N
concentration of leaves, and associated properties, e.g. chlorophyll content, are less
likely to have large resolution power than methods that assess nitrate-N of stem ends
or petioles, as was confirmed in Wu et al. (2007). This line of reasoning would
identify vegetation indices, derived from reflectance patterns in the visible and near
infrared region, as promising methods.

Environmental Aspects of Fertiliser Nitrogen Supply

Monotonous decline in ANR, even in the range of supplies where N remains limiting
production, corresponds to a yield response curve with monotonous decline in
marginal yield benefit. In northwestern Europe, the economical optimal point of
input is commonly in the range of 150–250 kg ha−1. At those levels of input, ANR
commonly drops to values of 0.5 to 0.6. ANR=0.6 means that 60% of the nitrogen
added to the system is resident in the crop biomass at harvest. Since the harvest
index for nitrogen is typically 0.75 (Vos 1997) and since only tubers leave the field,
the percentage of the added nitrogen remaining in the soil system is 55%, i.e. more
than half. In line with this, the amount of residual soil mineral N at harvest is
commonly larger after potato than after cereals or sugar beet (Neeteson 1994; Vos
and van der Putten 2000). Part of the nitrogen given in excess to offtake with tubers
carries over to the succeeding crops. However, part of it is lost over winter when (in
Western Europe) rainfall exceeds evapotranspiration, resulting in washing out of
nitrate on lighter textured soils and in waterlogging and lack of oxygen in fine
textures soil profiles, i.e. conditions inductive for denitrification. In a cropping
system on sandy soil, comprising spring wheat, sugar beet, oats and potato, Vos and
van der Putten (2004), corroborating findings of Shepherd (1999), observed that
systematic cultivation of catch crops after each main crop reduced the nitrate
concentration (averaged over the rotation and the winter season) in leachate at 80-cm
depth below the drinking water standard, whereas this was not the case without catch
crops. When potato is grown at economically optimum levels of N input (i.e. when
applying official recommendations), it is unlikely that the upper soil water of
sandy fields will show a seasonal averaged nitrate-nitrogen concentration less
than 50 mg l−1 nitrate, let alone 25 mg l−1 which is seen as the longer term
standard for groundwater nitrate concentration. Therefore, it is important that
farmers are allowed to compensate a ‘polluting’ crop with non-polluting crops in a
crop rotation.

Main Remaining Problems to Be Solved in Nitrogen Management

In the economically relevant range of N supply potato is a crop with a relatively low
ANR (about 0.5). The excess of nitrogen is bound to result in emissions to the
environment. As indicated, the drinking water standard for nitrate is easily exceeded
in leachate from light-textured soils. More stringent norms will increase the pressure
on the potato industry to manage crop nutrition as best as is possible. The concrete
316 Potato Research (2009) 52:305–317

impact of such more stringent norms on the nutrient input (and by consequence on
potato production) depends on a number of issues. These include:
(a) The actual value of the norms and the depth in the groundwater at which
compliance with the nitrate norms is measured.
(b) Whether or not surface water norms also apply to ditches between fields or
only to waters leaving agricultural areas.
(c) Lastly, the degree of coverage of the crop demand by organic fertilisers. This is
important since organic fertilisers commonly show lower ANR than chemical
fertilisers.
The implementation of the Water Framework Directive will tend to reduce
nutrient input in potato production. Hence, maximum utilisation of available
nutrients assumes higher importance. Precision agriculture (PA) and site-specific
management (SSM) will probably contribute to achieving a higher degree of nutrient
utilisation and hence help to comply with environmental norms. This requires further
development of a diagnostic system of collection of data from soil and crop that
leads to correct diagnosis of the nutrient status of the crop, distinguishing nitrogen
deficiency from other causes of spatially divergent crop performance. After a slow
start, PA and SSM seem to have caught the interest of farmers, which means that the
technology will probably diffuse into farming practice.

References

Abrahamsen P, Hansen S (2000) Daisy: an open soil–crop–atmosphere system model. Envir Modelling
and Software 15:313–330
Biemond H, Vos J (1992) Effects of nitrogen on the development and growth of the potato plant. 2. The
partitioning of dry matter, nitrogen and nitrate. Ann Bot 70:37–45
Breimer T (1989) Stikstofbijmestsysteem (NBS) voor enige vollegrondsgroenten. Consulentschap voor
Bodem- Water-, en Bemestingszaken in de Akkerbouw en Tuinbouw, Wageningen
Colnenne C, Meynard JM, Roche R, Reau R (2002) Effects of nitrogen deficiencies on autumnal growth
of oilseed rape. Eur J Agron 17:11–28
Ducheyne S, Schadeck N, Vanongeval L, Vandendriessche H, Feyen J (2001) Assessment of the
parameters of a mechanistic soil–crop–nitrogen simulation model using historic data of experimental
field sites in Belgium. Agric Water Manag 51:53–78
Gastal F, Lemaire G (2002) N uptake and distribution in crops: an agronomical and ecophysiological
perspective. J Exp Bot 53:789–799
Geijpens M, Vandendriessche H (1996) Advisory systems for nitrogen fertilizer recommendations. Plant
Soil 181:31–38
Gianquinto G, Goffart JP, Olivier MM, Guarda G, Colauzzi M, Dalla Costa L, Delle Vedove G, Vos J,
MacKerron DKL (2004/2005) The use of hand-held chlorophyll meters as a tool to assess the nitrogen
status and to guide nitrogen fertilization of potato crop. Potato Res 47:35–80
Goffart JP, Olivier M, Destain JP (2005) Presentation of a decision-support system (DSS) for nitrogen
management in potato production to improve the use of resources. In: Haverkort AJ, Struik PC (eds) Potato
in progress: science meets practice. Wageningen Academic Publishers, Wageningen, pp 134–142
Greenwood DJ, Draycott A (1988) Recovery of fertilizer-N by diverse vegetable crops: processes and
models. In: Jenkinson DS, Smith KA (eds) Nitrogen efficiency in agricultural soils. Elsevier Applied
Science, London, pp 46–61
Greenwood DJ, Lemaire G, Gosse G, Cruz P, Draycott A, Neeteson JJ (1990) Decline in percentage N of
C3 and C4 crops with increasing plant mass. Ann Bot 66:425–436
Jongschaap REE (2006) Run-time calibration of simulation models by integrating remote sensing
estimates of leaf area index and canopy nitrogen. Eur J Agron 24:316–324
Potato Research (2009) 52:305–317 317

Jongschaap REE, Booij R (2004) Spectral measurements at different spatial scales in potato: relating leaf,
plant and canopy nitrogen status. Int J Appl Earth Observation and Geoinformation 5:205–218
Liebscher G (1895) Untersuchungen über die Bestimmung des Düngerbedürfnisses der Ackerböden und
Kulturpflanzen. J Landwirtschaft 43:49
Lorenz H-J, Schlaghecken J, Engl G (1985) Gezielte Stickstoffversorgung. Das kulturbegleitende Nmin
Sollwerte-System (KNS-System). Deutscher Gartenbau 13:646–648
Marshall B, Vos J (1991) The relation between the nitrogen concentration and photosynthetic capacity of
potato (Solanum tuberosum L.) leaves. Ann Bot 68:33–39
Mitscherlich E (1909) Das Gesetz des Minimum, das Gesetz des Abnehmenden Bodenertrages.
Landwirtschaftliches Jahrbuch der Schweiz 38:537–552
Neeteson JJ (1994) Residual soil nitrate after application of nitrogen fertilizers to crops. In: Adriano DC,
Iskandar AK, Murarka IP (eds) Groundwater contamination. Advances in environmental science.
Science Reviews, Northwood, pp 347–365
Neeteson JJ, Wadman WP (1987) Assessment of economically optimum application rates of fertilizer N on
the basis of response curves. Fertiliser Res 12:37–52
Neeteson JJ, Greenwood DJ, Draycott A (1987) A dynamic model to predict yield and optimum nitrogen
fertiliser application rate for potatoes. Proceedings no. 262. The Fertiliser Society, London
Nitrate Directive (1991) Council Directive 91/676/EEC of 12 December 1991 concerning the protection of
waters against pollution caused by nitrates from agricultural sources. Official Journal L 375, 31/12/1991:1–8
Nitsch A, Varis E (1991) Nitrate estimates using the Nitracheck test for precise N-fertilization during plant
growth and, after harvest, for quality testing of potato tubers. Potato Res 34:95–105
Rémy J-C (1981) Etat actuel et perspectives de la mise en oeuvre de techniques de prévision de la fumure
azotée. CR Acad Agr Fr 67:859–874
Shepherd MA (1999) The effectiveness of cover crops during eight years of a UK sandland rotation. Soil
Use Management 15:41–48
van Alphen J (2002) A case study on precision nitrogen management in Dutch arable farming. Nutr Cycl
Agroecosystems 62:151–161
van der Ploeg RR, Bohm W, Kirkham MB (1999) On the origin of the theory of mineral nutrition of plants
and the law of the minimum. Soil Sci Soc Am J 63:1055–1062
van Keulen H (1982) Graphical analysis of annual crop response to fertiliser application. Agric Systems 9:113–126
Von Liebig J (1840) Die organische Chemie in ihrer Anwendung auf Agrikultur und Physiologie.
Friedrich Vieweg und Sohn Publ Co, Braunschweig
Vos J (1992) A case history: hundred years of potato production in Europe with special reference to The
Netherlands. Amer Potato J 69:731–751
Vos J (1997) The nitrogen response of potato (Solanum tuberosum L.) in the field: nitrogen uptake and
yield, harvest index and nitrogen concentration. Potato Res 40:237–248
Vos J (1999) Split nitrogen application in potato: effects on accumulation of nitrogen and dry matter in the
crop and on the soil nitrogen budget. J Agric Sci, Camb 133:263–274
Vos J, Biemond H (1992) Effects of nitrogen on the development and growth of the potato plant. 1. Leaf
appearance, expansion growth, life spans of leaves and stem branching. Ann Bot 70:27–35
Vos J, van der Putten PEL (1998) Effect of nitrogen supply on leaf growth, leaf nitrogen economy and
photosynthetic capacity in potato. Field Crops Res 59:63–72
Vos J, van der Putten PEL (2000) Nutrient cycling in a cropping system with potato, spring wheat, sugar
beet, oats and nitrogen catch crops. I. Input and offtake of nitrogen, phosphorus and potassium. Nutr
Cycl Agroecosystems 56:87–97
Vos J, van der Putten PEL (2001) Effects of partial shading of the potato plant on photosynthesis of treated
leaves, leaf area expansion and allocation of nitrogen and dry matter in component plant parts. Eur J
Agron 14:209–220
Vos J, van der Putten PEL (2004) Nutrient cycling in a cropping system with potato, spring wheat, sugar
beet, oats and nitrogen catch crops. II. Effect of catch crops on nitrate leaching in autumn and winter.
Nutri Cycl Agroecosystems 70:23–31
Vos J, van der Putten PEL, Birch CJ (2005) Effect of nitrogen supply on leaf appearance, leaf growth, leaf
nitrogen economy and photosynthetic capacity in maize (Zea mays L.). Field Crops Res 93:64–73
Water Framework Directive (2000) Directive 2000/60/EC of the European parliament and of the Council
of 23 October 2000 establishing a framework for community action in the field of water policy. Off J
Eur Communities L327:1–72
Wu J, Wang D, Rosen CJ, Bauer ME (2007) Comparison of petiole nitrate concentrations, SPAD
chlorophyll readings, and QuickBird satellite imagery in detecting nitrogen status of potato canopies.
Field Crops Res 101:96–103