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SEASONAL FLUCTUATION AND INTRACANOPY VARIATION

IN LEAF NITROGEN LEVEL IN OLIVE
a
S. Perica
a
Institute for Adriatic Crops, Put Duilova 11, Split, 21000, Croatia
Published online: 16 Aug 2006.

To cite this article: S. Perica (2001): SEASONAL FLUCTUATION AND INTRACANOPY VARIATION IN LEAF NITROGEN LEVEL IN
OLIVE, Journal of Plant Nutrition, 24:4-5, 779-787

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 JOURNAL OF PLANT NUTRITION, 24(4&5), 779±787 (2001)

SEASONAL FLUCTUATION AND

INTRACANOPY VARIATION IN LEAF
NITROGEN LEVEL IN OLIVE
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S. Perica

Institute for Adriatic Crops, Put Duilova 11, 21000 Split,

Croatia

ABSTRACT

Seasonal ¯uctuation and intracanopy variation in leaf nitrogen

(N) of bearing olive (Olea europaea L.) was assessed. The
N concentration was high and stable during the wintertime
(about 1.8% dwt). With the advancement of vegetative growth
and reproductive development, N concentration in the leaves
decreased sharply, reaching the lowest value (about 1.45 to
1.50% dwt) in summer months June, July, and early August,
coincident with early fruit development. Among intracanopy fac-
tors studied, the largest single in¯uence on N concentration was
light exposure. Thus on average, over all leaf types and canopy
heights, leaves from the south side of the tree had 11.8% more
N than north side leaves. Consistencies with regarding sampling
time, canopy locations, height, and nodal position of the leaf are
critical for diagnosis of N status in the olive.

INTRODUCTION

Nitrogen is perhaps the most important mineral element for olive

production in most of olive growing regions. It has been shown that N levels

779

Copyright # 2001 by Marcel Dekker, Inc. www.dekker.com

 780 PERICA

directly affect fruit set, yield, and shoot growth in the olive (1, 2, 3, 4). Leaf
analysis is widely used to monitor the olive N level and help plan fertilizer
programs (5, 6, 7).
Variations in nutrient concentrations during the season or in tissues of
different age have been reported for a number of crops. Although there are such
studies with bearing trees for apples and other crops (8), in case of olives
frequently young plants have been used, often grown in sand or solution cultures
(9, 10, 11, 12, 13). While, this avoids the variability problems associated with
sampling larger plants, it is unlikely that results from these studies are relevant to
mature trees. When mature plants have been used, differences found in leaf N
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concentration commonly have been attributed to individual tree productivity,

different cultivars or environments (14, 15). Although each of these factors
clearly in¯uence leaf N values, there is a need to determine within-tree N
variability so that appropriate sampling techniques can be developed.
Jones (16) emphasized that the method of leaf collection can have a
profound effect on the results obtained and that no other component of the plant
analysis technique can have as much effect on the ®nal result and its
interpretation. This particularly applies to olive where the life span of olive
leaves is of the order of 2±3 years, and a single olive shoot can have leaves of
distinctly different ages and physiological activities.
The objective of this study was to assess the seasonal ¯uctuation in N
concentration in the leaves of bearing olive tree, to identify possible sources of
that variability and to determine its magnitudes. Understanding the nature of leaf
N variability could assist further development of leaf analysis technique in the
olive. It was of particular interest, therefore to evaluate the in¯uence of canopy
and nodal position, and time of sampling on N levels in the olive leaves.

MATERIALS AND METHODS

Twelve years after rejuvenation from the trunk, ten olive trees cv. ``Oblica'',
were selected on an Adriatic island (Latitude: 43 210 N, Longitude: 16 330 E) for
regular monthly leaf samplings. Similar ®ve trees were selected on another nearby
site for samplings within the canopy. All the trees were trained into a globe-like
shape, medium bearing, dry farmed, uniform in size, and yield. The orchard rows
were oriented east-west at 2.567 m spacing with a canopy height of 3.5 m.
For determination of seasonal effect on leaf N concentration, 100 fully
developed leaves from two adjacent trees (total of ®ve replications per sampling)
were collected monthly from the central part of the shoots. Sampled shoots were
equally distributed all around the canopy at height of about 1.8 m. Extremely
vigorous or weak shoots were avoided in all samplings. Various tree development
stages were also recorded.
 LEAF NITROGEN LEVEL IN OLIVE 781

For the effect of canopy position on leaf N level, leaf samples were taken in
winter (January). Winter time in this area lasts about four to ®ve months, and is
considered to be the minimal ¯ux period for olive leaf nutrients, thus suitable for
leaf sampling (5). Three main factors considered were: canopy light exposure
(E, S, W, and N side), canopy height (lower section 1.1±1.4 m; central 1.4±1.7 m;
high section 1.7±2.1 m), and relative leaf nodal position (basal, central, and apical
leaf). Average shoots were 15.28 cm long with 7.75 nodes per shoot, on such a
shoot the ®rst one or two nodes and the last one or two nodes would not be
sampled. From each of ®ve trees, 36 samples of 20 leaves each were taken per
canopy. The experiment was considered split±split plot design with canopy side
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as a main plot, canopy height as a subplot, and leaf nodal position as sub-sub plot.
In the collected samples leaf N was determined by Kjeldahl digestion using
Kjeltec System 1026 (17).

RESULTS AND DISCUSSION

Time of Sampling

The N concentration was high and stable during the wintertime, from
October to March (Figure 1). Beginning from April, concentration of N in the
leaves started to decrease sharply. The difference coincides with bud break and is
due to onset of vegetative growth and initiation of reproductive activities. With
the advancement of vegetative growth and reproductive development, N
concentration decreased reaching the value of 1.53 to 1.65% dwt at time of
¯owering and fruitset. The lowest values were found in summer months June,
July and early of August coincident with early fruit development (1.45±
1.50% dwt).
Priestley (18) illustrated signi®cance of the olive leaves as important organs
for carbohydrate storage. Olive leaves also represent storage organs for N (19)
and they release N in response to the demand of various metabolic sinks. During
periods of high N demand and an inadequate storage of N in other pools, leaves
may supply an increased proportion of N requirement. This explains our
observation of high and stable N levels during winter, at time of late fruit
maturation and practical cessation of vegetative growth when a minimal demand
for N could be expected. Furthermore, low levels of N in spring and in summer
time, coincide with the time of strong demand for N in developing reproductive
and vegetative organs and presumably reduced soil supply before autumn rains
occur.
The stability of N in the olive leaves in winter, and similar stability during
early summer, were used to propose these two periods as the most appropriate for
the assessment of nutrient status of the olive. Our data indicate that in summer
 782 PERICA
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Figure 1. Changes in nitrogen concentration and total content in the leaves of bearing
olive tree during an annual growth cycle. Fully developed leaves were collected from the
central part of the shoots. Sampled shoots were equally distributed all around the canopy at
height of about 1.8 m. Error bars represent one standard error of the mean of ®ve
replicates.

under given experimental conditions the stable concentration is obtainable for

about two months, which is somewhat shorter than in winter, when the
concentration of N is steady over three or four months. Suggested norms for the
interpretation of nutrient status, mostly compiled from early works, are: a) for a
winter sample: optimum 1.6±1.8%; high 1.8±2.2% (20); critical level 1.5%,
optimum 2.1% (5, 21), b) for summer sample: adequate 1.5±2.0% (6), and also
adequate 1.5±2.5% (22). Comparing values obtained from this study with the
listed values it can be concluded that the experimental trees were probably sub-
optimally supplied by N in summer time. While the same trees in winter could be
 LEAF NITROGEN LEVEL IN OLIVE 783

described as having optimum N. Although this work did not aim to alter current
recommended leaf nutrient concentrations, since it did not take into account any
year-to-year variations, yield status or yield-nutrient relationship, it still indicates
that the winter level of stored N in the leaves could be important since N is
exported during vegetative onset and reproductive development. Thus, under the
presumption that stored N is important then further determination of the optimum
level of N in winter leaf sample should be examined.

Canopy and Nodal Position

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Canopy and nodal position in¯uenced leaf N concentration (Table 1).

Higher concentration of N was found in the leaves from south side of the canopy,
in the leaves from higher portion of the canopy, and in the younger leaves.
Among the factors studied, the largest single in¯uence on N concentration was
the canopy side. Thus on average, over all leaf types and canopy heights, leaves
from the south side of the tree had 11.8% more N than north side leaves
(P < 0.001). These differences became more apparent when in addition to canopy
side, different canopy heights, and leaf ages are compared (Figure 2). This
phenomenon is believed to be related to photosynthesis ef®ciency and N
consumption. Changing radiation environment, given a ®xed amount of N
available to leaves, plants allocate N in order to optimize total whole canopy
photosynthesis (23).
Leaf analysis can be a powerful tool in diagnosing N status of the tree but
samples must be taken carefully to insure that valid comparison with reported
standards can be made. In addition to the factors studied potential differences
in result due to other commonly accepted causes of variability such as soil,
environment, cultivar fertilizer application must also be considered. Data from
this study are assumed to be representative for wider range of conditions,

Table 1. The Main Effect of Canopy and Nodal Leaf Position

on Olive Leaf Nitrogen Concentration (N% DM)

Main Factors

Canopy Side Canopy Height Nodal Position

South 1.89 a High 1.86 a Basal 1.84 a
East 1.87 b Central 1.80 b Central 1.81 b
West 1.79 b Low 1.78 b Apical 1.79 b
North 1.70 c

Means in a column followed by the same letter are not

signi®cantly different (P < 0.05).
 784 PERICA
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Figure 2. Concentration of nitrogen in the winter olive leaf samples in relation to aspect
(E, S, W, N) canopy heights (low, central, high) and relative nodal position of the leaf
(basal, central and apical). Error bars represent one standard error of the mean of ®ve
replicates.
 LEAF NITROGEN LEVEL IN OLIVE 785

however training system, may show different magnitude of within three variation
due to changing pattern of light penetration, which may be affected by planting
density, tree size, or climate.

CONCLUSIONS

Consistency with regarding canopy side, height, nodal position of the leaf,
and sampling time of the year is critical for diagnosis of N status in the olive.
Current standards are based on the most recent mature summer or winter leaf.
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However our data show that, at least in winter, relatively large variations in leaf
concentration could result from changing the side of the canopy, canopy height or
even sampling position on the shoot. Comparing the N level of such a leaf with
the proposed standard may lead to incorrect conclusions. Results from this study
indicate that variation can be minimized by good sampling protocol, which
involves sampling each tree uniformly all around the canopy, sampling at similar
height and sampling adequate number of trees.

ACKNOWLEDGMENTS

The author gratefully acknowledges I. Skejic and N. Goreta for technical

assistance, Dr. P. H. Brown and Dr. H. Hu for critical review of this manuscript,
and an anonymous reviewer for many valuable suggestions.

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