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Seasonal Changes of Mineral Nutrients in Olive Leaves During The Alternate-Bearing Cycle
Seasonal Changes of Mineral Nutrients in Olive Leaves During The Alternate-Bearing Cycle
Abstract
Samples of olive (Olea europaea L.) leaves developed in 1993, 1994 or 1995 were collected
separately from non-irrigated, mature trees at monthly intervals during 1994 (`off' year) and 1995
(`on' year) to determine leaf-nutrient concentration and nutrient content per leaf. N, P, K and Mg
contents were affected by crop load, showing lower values following the `on' year. No effect of crop
load was observed on micronutrient accumulation in leaves, probably because of the lower
requirements for these elements of the olive fruit. Leaf age also influenced leaf-nutrient
concentration and nutrient content. Leaf N, P, K, Zn and B concentrations were higher in younger
leaves, whereas leaf Ca, Mg, Mn, Cu and Fe concentrations were higher in older leaves. Ca, Mg,
Mn, Cu and Fe content per leaf were also higher in older leaves. There was little or no difference in
P, K, Zn and B content among leaves of different ages. N content per leaf was higher in older leaves
during the growing period of the `off' year. Seasonal mineral fluctuations curves for different leaf
mineral nutrients were developed which could be used in the interpretation of leaf analysis. # 1999
Elsevier Science B.V. All rights reserved.
1. Introduction
The olive (Olea europaea L.) is an evergreen tree that has been cultivated from
ancient times. Presently, there are more than eight million hectares producing
0304-4238/99/$ ± see front matter # 1999 Elsevier Science B.V. All rights reserved.
PII: S 0 3 0 4 - 4 2 3 8 ( 9 9 ) 0 0 0 4 5 - X
26 R. FernaÂndez-Escobar et al. / Scientia Horticulturae 82 (1999) 25±45
olives in the world, most of them located in the Mediterranean Basin. Olives
flower on one-year-old wood and display two inflorescences per node. In the
northern hemisphere, flower bud induction is manifested by July, around the time
of endocarp sclerification (FernaÂndez-Escobar et al., 1992). Floral differentiation
is manifested by March (Hartmann, 1951), and anthesis occurs by May (AlcalaÂ
and Barranco, 1992). Shortly after anthesis, massive abscission of flowers and
fruits occurs (Rallo and FernaÂndez-Escobar, 1985). The remaining fruits usually
persist on the tree until harvest, which takes place during the fall and winter.
The olive is strongly alternate-bearing. This phenomenon may influence
nutrient content of the trees and the annual nutrient consumption, as has been
reported for pistachio (Brown et al., 1995), another alternate-bearing tree crop
species.
Leaf-nutrient analysis is the best method for diagnosing tree nutritional status,
and represents an important tool for determining future fertilization requirements
(Benton Jones, 1985). Presently, the use of leaf analysis as a guide for olive
fertilization is still infrequent in Spain and in other Mediterranean countries,
although the technique is becoming popular. Lack of dissemination of this
technique may be due, in part, to the lack of available information on nutritional
requirements of olive trees and also on the limitations of leaf analysis as a tool for
guiding fertilization programs. In addition, there has been confusion regarding the
appropriate time for leaf collection, a critical aspect of this technique.
It is well known that leaf analysis must be based on standardized sampling
methods and that results must be compared only with standard values obtained by
those procedures. Standard values for olives have been compiled by FernaÂndez-
Escobar (1997) based on data reported by Chapman (1966), Childers (1966) and
Beutel et al. (1983). These values refer to nutrient concentrations in recently
mature leaves sampled in July in the northern hemisphere from the middle
portion of non-bearing, current-season shoots. However, the seasonal variations in
leaf-nutrient concentration and leaf-nutrient content are necessary in order to
understand the physiological aspects of olive nutrition, and are also helpful in the
interpretation of leaf analysis. These seasonal changes are not available for olives.
The determination of such changes during the alternate-bearing cycle was the
objective of the present work.
intervals during 1994 (`off' year) and 1995 (`on' year) from four blocks of 50
olive trees each. Two or three fully-expanded, mature leaves per experimental tree
were collected each time from the current season growth of 1994 or 1995, and
also from the 1993 growth in 1994 and 1995, and from the 1994 growth in 1995.
In all cases, leaves were taken from the middle portion of the shoot corresponding
to each growth period. Leaf samples of the same age from each block were
combined as samples of 100±150 leaves for the determination of nutrient
concentration and nutrient content per leaf.
Leaves were collected in paper bags and stored in a portable ice chest. Once in
the laboratory, leaves were washed with 0.03% Triton X-100, rinsed in deionized
water, dried at 808C for 48 h, ground, and stored in an oven at 608C until analysis.
Nitrogen was determined by the Kjeldahl procedure. For other element
determinations, the stored samples were ashed in a muffle furnace at 6008C for
12 h, and dissolved in 0.1 N HCl. Total P was determined by colorimetry using
the method described by Murphy and Riley (1962). Boron was determined in the
extract by colorimetry (Greweling, 1976). The remaining elements (K, Mg, Ca,
Zn, Mn, Fe, and Cu) were measured using an atomic absorption spectro-
photometer Perkin±Elmer 1.100 B. Mean and standard errors of each sample
were calculated for statistical comparison.
3.2. Nitrogen
Leaf N concentration was higher in younger leaves in both, `off' and `on' years
(Fig. 2). Nitrogen concentration of current-season leaves of 1994 and 1995 began
to drop in June, reaching a minimum value in August. Subsequently, during
autumn, these leaves showed an increase in N concentration reaching a maximum
at the end of the season. These values were almost constant until the following
June in leaves of the 1994 growth, when a new drop in N concentration was
28 R. FernaÂndez-Escobar et al. / Scientia Horticulturae 82 (1999) 25±45
Fig. 1. Seasonal changes in the dry mass of fully-expanded olive leaves developed during three
different growth periods and an alternate-bearing cycle.
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30 R. FernaÂndez-Escobar et al. / Scientia Horticulturae 82 (1999) 25±45
3.3. Phosphorus
Leaf P concentration was higher in younger leaves in both, the `off' and `on'
years (Fig. 3). Phosphorus concentration of the current-season leaves of
1994 dropped in June to reach a minimum in September. Subsequently, during
autumn, these leaves increased in P concentration, to reach a maximum in the
winter, showing a new drop in P concentration in March 1995 and decreasing the
same year. Leaves of the 1993 growth showed a pattern similar to leaves
developed in 1994, but they reached lower concentrations. Phosphorus
concentration of leaves from 1995 current-season growth also showed a pattern
similar to that of the 1994 growth, dropping to a minimum in August and
increasing during autumn, with a subsequent drop in October. Leaf P con-
centration of July samples of the current-season growth of both, 1994 and 1995
showed values above the threshold limit for sufficiency of 0.1%, but was higher in
the `on' year.
Phosphorus content of the three kinds of leaf samples showed similar
patterns of P concentration. Phosphorus accumulated in all the leaves during
the autumn and winter following the `off' year, and then decreased during
the `on' year. There were less pronounced differences in leaf P content
among leaf ages than that observed in leaf P concentration. The general dec-
rease in leaf P content during the `on' year could be due to P consumption by the
fruits.
The seasonal changes in leaf P concentration and content per leaf are generally
in agreement with those reported for other tree fruit species. Leaf P concentration
decreased throughout the growing season in deciduous trees (Ryugo, 1988; Shear
and Faust, 1980), citrus (Embleton et al., 1973) and avocado (Embleton and
Jones, 1966). We observed the same trends in olive trees, and also an increase in
leaf P concentration during the autumn in both years. This tendency has not been
reported consistently in other evergreen trees, although fluctuations in leaf P
concentration were observed by Bingham (1961) in avocado and by Fahmy and
Nasrallah (1959) in olive. However, the data reported by the latter authors are in
contrast with the results presented here. They found the lowest leaf P
concentration values in April and May and the highest in January in one year
and in August the next year, whereas we always found a minimum in August or
September and an accumulation of P in leaves during the autumn and winter,
following the `off' year. Accumulation of P in leaves following the `off' year was
also observed by Brown et al. (1995) in pistachio. The sample procedure adopted
in the present work, where leaves of three different ages were collected
separately, better reflects the seasonal trends of mineral nutrients in leaves and
permits a more precise interpretation of the data. The differences in sampling
methodology may explain, at least in part, the disagreement of our data with those
reported by Fahmy and Nasrallah (1959).
R. FernaÂndez-Escobar et al. / Scientia Horticulturae 82 (1999) 25±45
Fig. 3. Seasonal trends of P concentration and content in olive leaves developed during three different growth periods and an alternate-bearing cycle.
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3.4. Potassium
3.5. Calcium
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34
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Fig. 5. Seasonal trends of Ca concentration and content in olive leaves developed during three different growth periods and an alternate-bearing cycle.
R. FernaÂndez-Escobar et al. / Scientia Horticulturae 82 (1999) 25±45 35
3.6. Magnesium
3.7. Manganese
Leaf Mn concentration was usually higher in the older leaves of both, the `off'
and `on' years (Fig. 7). Manganese concentration of the current-season growth
leaves of both the years increased from the beginning of the season, reaching a
maximum in July. Subsequently, Mn concentration in leaves decreased through
the end of the season. In leaves developed in 1994, there was a slight increase in
Mn concentration during the `on' year of 1995. Leaves developed in 1993 showed
a decrease in Mn concentration during the `off' year, and then tended to increase
it in the `on' year. Leaf Mn concentration of the July samples of current-season
growth in both, the `off' and `on' years were above the threshold limit for
sufficiency of 20 mg kgÿ1, but it was lower in the `on' year.
Leaf Mn content was significantly higher in older leaves. Manganese content of
leaves developed in 1993 decreased during the `off' year, and then increased
36
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Fig. 6. Seasonal trends of Mg concentration and content in olive leaves developed during three different growth periods and an alternate-bearing cycle.
R. FernaÂndez-Escobar et al. / Scientia Horticulturae 82 (1999) 25±45
Fig. 7. Seasonal trends of Mn concentration and content in olive leaves developed during three different growth periods and an alternate-bearing cycle.
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38 R. FernaÂndez-Escobar et al. / Scientia Horticulturae 82 (1999) 25±45
slightly in the `on' year. In contrast, leaves developed in 1994 showed a slight Mn
accumulation during 1994 and 1995, and leaves from the 1995 growth had
noticeable fluctuations in Mn content but were constant after July. Little
differences were observed between `off' and `on' years in Mn content per leaf,
although it was slightly higher following the `off' year.
Leaf Mn concentration tends to increase with time in deciduous fruit trees
(Brown, 1994; Buwalda and Meekings, 1990; Shear and Faust, 1980), but the
reverse has been reported in some evergreen species (Broschat, 1997). In the
olive, the maximal Mn concentration was measured in July in leaves from
current-season growth. Increases of leaf Mn concentration were observed in older
leaves during the `on' year, but not during the `off' year.
3.8. Zinc
Leaf Zn concentration was higher in younger leaves in both `off' and `on' years
(Fig. 8). Zinc concentration of current-season growth leaves of both years clearly
decreased throughout the season to a minimum in December. Leaves from the
1994 growth remained at these levels during the next year. Leaves developed in
1993 showed little fluctuation during both these years, with a small peak in May
1994. Leaf Zn concentration of July samples of current-season growth in both
these years were above the threshold limit for sufficiency of 10 mg kgÿ1, and was
lower in the `on' year.
Leaf Zn content showed little differences between the three kinds of leaves
sampled. Zn accumulated in leaves following the `off' year to a maximum in
April 1995, and diminished during the `on' year of 1995. During the `off' year
there was a maximum in May±June in both kinds of leaves.
Little information is available concerning Zn nutrition of olives. However, the
seasonal trends presented in Fig. 7 are typical for this element in deciduous tree
species (Ryugo, 1988; Shear and Faust, 1980). In the olive, little difference was
found between `off' and `on' years.
3.9. Copper
Leaf Cu concentration was higher in leaves developed in 1993 than in the other
leaves, and no differences were found between leaves developed in 1994 and
1995 (Fig. 9). Leaves from 1993 growth showed large fluctuations but no clear
seasonal trend. In contrast, leaves from the 1994 and 1995 growth seasons
maintained its values almost constants during the season. Leaves developed in
1994 exhibited a slight increase in Cu concentration throughout 1995. Leaf Cu
concentration of July samples of the current-season growth of both these years
were above the threshold limit for sufficiency of 4 mg kgÿ1. Leaf Cu content
R. FernaÂndez-Escobar et al. / Scientia Horticulturae 82 (1999) 25±45
Fig. 8. Seasonal trends of Zn concentration and content in olive leaves developed during three different growth periods and an alternate-bearing cycle.
39
40
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Fig. 9. Seasonal trends of Cu concentration and content in olive leaves developed during three different growth periods and an alternate-bearing cycle.
R. FernaÂndez-Escobar et al. / Scientia Horticulturae 82 (1999) 25±45 41
showed the same trend that Cu concentration had in all leaves. However, leaf Cu
content was generally higher in older leaves.
There are no reported cases of Cu deficiency in olives, mainly because copper
is used as a fungicide to control the olive leaf spot caused by Spilocea oleagina
(Castagne) Hughes. Periodic Cu sprays explain the great accumulation of this
element, particularly in the oldest leaves, and the erratic fluctuations observed in
leaf Cu concentration and content per leaf. Seasonal trends of Cu concentration of
the current-season growth leaves are, moreover, similar to those reported for other
tree fruits (Brown, 1994).
3.10. Boron
Leaf B concentration was higher in younger leaves in both years (Fig. 10).
Boron concentration of the current-season growth leaves of both, the `off' and
`on' years showed a general decrease throughout the season, but depleted in May
or June. This depletion occurs around anthesis and has been linked to B
mobilization from the leaves to supply B requirements of flowers and fruits
(Delgado et al., 1994). Leaves from the 1993 growth showed little fluctuations in
B concentration throughout the `off' year, and tended to increase it in 1995. Leaf
B concentration of July samples of the current-season growth in both these years
were within the adequate range of 19±150 mg kgÿ1, showing no differences
between years.
There was little difference in leaf B content among the three kinds of leaves
sampled. Leaf B content per leaf increased until June-July within each year, and
then decreased until October±November. Little difference was observed between
the `off' and `on' years, although in the `on' year the maximum B content per leaf
was higher than in the `off' year.
Seasonal trends in leaf B concentration varies greatly among species. Leaf B
concentration increases in citrus (Embleton et al., 1973) and fig (Brown, 1994) as
the season progresses, but in avocado (Embleton and Jones, 1966) and Japanese
pear (Buwalda and Meekings, 1990) leaf B decreases, as we observed in olive.
Moreover, olive require high levels of boron (Hartmann et al., 1966) and,
therefore, B fertilizer is applied in many olive orchards.
3.11. Iron
Only data from the `off' year of 1994 are available for iron. During this year,
leaf Fe concentration and content per leaf were higher in older leaves (Fig. 11).
Leaf Fe concentration fluctuated throughout the season, showing a general
tendency to oscillate around a value of 40 mg kgÿ1 for one-year-old leaves and
20 mg kgÿ1 for the current-season leaves. A similar pattern has been observed in
other tree species (Embleton and Jones, 1966; Buwalda and Meekings, 1990;
42
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Fig. 10. Seasonal trends of B concentration and content in olive leaves developed during three different growth periods and an alternate-bearing cycle.
R. FernaÂndez-Escobar et al. / Scientia Horticulturae 82 (1999) 25±45
Fig. 11. Seasonal trends of Fe concentration and content in olive leaves developed during two different growth periods.
43
44 R. FernaÂndez-Escobar et al. / Scientia Horticulturae 82 (1999) 25±45
Brown, 1994). Standard values have not been established for Fe in the olive. As
occurs for other species, leaf analysis is not useful for diagnosing Fe deficiency
because the inconsistency of leaf Fe levels in separating chlorotic from non-
chlorotic leaves (Korcak, 1987). Visual examination of trees is the best method
for diagnosing Fe deficiency, a nutritional problem that negatively affects olives
growing in alkaline, calcareous soils (FernaÂndez-Escobar et al., 1993).
4. Conclusions
Acknowledgements
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