1999.
Revista Chapingo Serie Horticultura 5:89-94.
THE ANNUAL
PRODUCTION AND UTILIZATION OF DRY MATTER OF AN AVOCADO (Persea
Gaaton, 25130 Israel. Fax.
972-4-9858413. E-mail: gadar@research.haifa.ac.il
This work, which was a part of the M.Sc. studies in the Faculty of
Agriculture in the Hebrew University of Jerusalem, was carried out under the
supervision of Prof. E.E. Goldschmidt. Two basic processes which take place
concurrently in the avocado tree during the yearly cycle of growth and
reproduction are discussed in this essay: Dry matter production and its
utilization.
Key Words: Tree physiology, reserves, productivity.
The production of dry matter and its distribution among plant organs are
basic processes on which other internal activities are dependent. Therefore it
is important to study the carbon balance of the avocado tree.
The objective of this essay is to
compile a dry matter budget for a fruiting avocado tree and to find its
relations to the yearly avocado growth cycle.
Eventually, this dry matter budget
may be part of a dynamic avocado productivity model, as developed for citrus by
Harpaz et al. (1990) and for other
crops. The data presented in the essay are based on photosynthetic measurements
made in
The data, calculations and
estimations presented in this paper, refer to an avocado tree which reaches a
canopy surface area of 50 m2 by the end of September. Such a tree
has been defined a “standard” tree.
The main data which are presented
in following sections are based on photosynthetic measurements made by Sharon
& Bravdo (unpublished) on the south quadrant of a ‘Hass’ avocado tree, on
the dry matter content of ‘Hass’ avocado tree tissues presented by Lovatt
(1996), on the avocado fruit growth presented by Ofer (1986) and on the yearly
growth of the perennial parts of an avocado tree which grows in a commercial
orchard.
Four rounds of photosynthetic
measurements, with an interval of two hours, conducted on September 21, 1977
(12 hours of daylight) on the external layer of the south quadrant of a ‘Hass’
avocado tree, revealed that the daily assimilation of a quadrate meter was 0.21
mol CO2. This figure refers to the hours of direct sunlight as well
as to indirect lighting during the day.
According to this figure, estimates of the
assimilation of the east, west and north quadrants have been made. So, it follows
that the daily assimilation rate of an avocado tree, with an external canopy
surface of 50 m2 is 5.18 of CO2.
According to data on reduced light
intensity which reaches leaves on the inner, shaded parts of the tree (Schaffer
and Andersen, 1994), it was estimated that for the whole tree, those leaves
assimilate 1.9 mol of CO2 per day. So, the total daily assimilation
of an avocado tree is 7.08 mol of CO2, which are 311.5 gram of CO2.
The impact of each of the
following five factors on the monthly assimilation rate of an avocado tree was
calculated: leaf age, day length, yearly canopy growth, eastern dry wind
stress, and climatic conditions during winter.
1.1)
The life span of an avocado leaf is approximately one
year.
1.2)
The flush of avocado leaves and their shedding occurs
in an uneven rate all year long (mostly between April and July).
1.3)
The maximum assimilation efficiency of avocado leaves
is attained between the 25th and 70th day after
sprouting. Afterwards, their assimilation efficiency declines gradually towards
senescence.
These are the physiological facts
according to which the term “assimilation potential” has been defined.
“Assimilation potential” is the
monthly assimilation yield, in mol CO2, of an avocado leaf,
calculated according to the order of the leaf flush and age. This factor refers
to the internal capability of the leaf to assimilate CO2.
Day length, eastern dry wind
stress and climatic conditions during winter, are external factors that have an
impact o the leaf’s assimilation rate.
As mentioned earlier, our estimate
of the assimilation of an avocado tree, 7 mol of CO2 per day, or 210
mol of CO2 per month, was calculated according to photosynthetic measurements
made on September 21, when the hours of daylight equal those of darkness.
During spring, the number of light
hours that extend by 5.5% per month while during autumn they are reduced at the
same rate. Therefore, the monthly assimilation potential has been adjusted
accordingly: the first adjustment.
It was estimated that the seasonal
growth of the canopy’s outer layer area of an avocado tree in a commercial
orchard reaches during summer, about 14%. Most of this yearly growth is
vertical. The second adjustment of monthly assimilation has been done according
to this estimation.
The reduction of the monthly
assimilation rate, caused by climatic conditions, such as spring east dry winds
and the winter climate, was the third adjustment. The third adjustment has been
done according to data provided by The Israeli Meteorological Service.
The figures of the calculated
monthly assimilation of an avocado tree, after the three adjustments mentioned
earlier, are presented in Table 1, line 2. According to the figures, the
highest monthly assimilation of 216 mol of CO2 is gained in August
by the “standard” avocado ‘Hass’ tree, while the lowest monthly assimilation
occur in February.
According to the figures, the
calculated monthly assimilation of an avocado tree in January and in February
are only about 29% of the potential monthly assimilation, while in August it is
about 97% (Table 1, line 2/1). The monthly gap between the calculated (actual) assimilation of an
avocado tree and its assimilation potential, demonstrates mostly the
influence of climatic conditions and canopy growth, on the avocado’s
assimilation performance.
|
Table 1.
The calculated monthly assimilation rate and the
assimilation potential of a “standard” avocado tree (mol of CO2
per month). |
||||||||||||
|
Month: |
A |
M |
J |
J |
Au |
S |
O |
N |
D |
J |
F |
Ma |
|
1 |
86 |
100 |
133 |
172 |
222 |
210 |
196 |
168 |
159 |
146 |
123 |
112 |
|
2 |
55 |
82 |
118 |
162 |
216 |
210 |
156 |
120 |
73 |
43 |
36 |
51 |
|
2/1 |
0.63 |
0.82 |
0.89 |
0.94 |
0.97 |
1.00 |
0.80 |
0.71 |
0.46 |
0.29 |
0.29 |
0.46 |
|
Month:
from April to March. 1: The
monthly assimilation potential of an avocado tree in mol CO2. 2: The
calculated monthly assimilation in mol of CO2 2/1: The
ratio of calculated monthly assimilation to the assimilation potential of a
“standard” avocado tree. |
||||||||||||
The yearly growth of an avocado tree
In order to evaluate the yearly
dry matter production, the following information was taken into account: Data
presented by Cameron (1952) and Lovatt (1996) about avocado trees that were
dismembered into organs. Their fresh and dry weights were reported; and growth
rates of avocado shoots and roots measured in a rhizotron by Ploetz et al. (1992).
The following assumptions were made for the compilation of Table 2:
· A crop of 53.6 kg, F.W. per tree
(1500 kg, F.W. per dunam).
· A yearly growth rate of 14% of
the perennial organs.
· A flesh/seed ratio of 84/16 in
the avocado fruit.
|
Table 2. The
yearly utilization of dry matter in the organs of an avocado tree. |
|||||
|
|
Fresh weight (g) |
Dry Matter (%) |
Dry Matter (g) |
Yearly Addition (%) |
Yearly addition of fresh weight (g) |
|
Perennial
Organs: |
176300 |
44.3 |
78100 |
14 |
10934 |
|
New
organs: Roots |
4100 |
37.1 |
1523 |
100 |
1523 |
|
Shoots |
1200 |
36 |
430 |
100 |
430 |
|
Leaves |
24000 |
40 |
9600 |
100 |
9600 |
|
Inflorescences |
8094 |
20.2 |
1643 |
100 |
1643 |
|
Total:
(Without fruit) |
213694 |
42.7 |
91296 |
26.4 |
24130 |
|
Fruit: |
53571 |
|
|
|
|
|
Fruit flesh (84%) |
45000 |
21 |
(9450) |
|
|
|
F.f.con. |
|
|
14660 |
|
|
|
Fruit Seed (16%) |
8571 |
40 |
3428 |
|
18088 |
|
Total:
(including fruit) |
267265 |
44.9 |
109384 |
38.6 |
42218 |
|
Perennial organs: scaffold roots, trunk and branches. FW (g): fresh weight,
grams. DW (g): dry matter, grams. F.f.con.: Oil and protein in the fruit flesh
converted to equivalent carbohydrate units. |
|||||
The yearly growth of every organ, presented in the Table 2, has been divided
to months according to data from research reports, or to the author’s
assumptions, when no data were available (Table 3).
|
Table 3.
The monthly increase in dry matter of an avocado tree
(total 42218). |
||||||||||||
|
Month: |
Ma |
A |
M |
J |
J |
Au |
S |
O |
N |
D |
J |
F |
WF
|
2147 |
1705 |
3507 |
4017 |
4003 |
3317 |
1713 |
1121 |
830 |
734 |
619 |
417 |
Fruit
|
|
|
123 |
261 |
1933 |
3663 |
5345 |
6763 |
|
|
|
|
Total
|
2147 |
1705 |
3630 |
4278 |
5936 |
6980 |
7058 |
7884 |
830 |
734 |
619 |
417 |
|
Month:
from March to February WF:
Without fruit. |
||||||||||||
The energy balance of the growing avocado fruit
The periodic accumulation of all
energy comprising components in an avocado fruit were calculated, while the fruit’s
own photosynthetic carbohydrate production was estimated for the same periods.
The calculations were based on the
periodical measurements of net efflux of CO2 of attached avocado
fruit cv. Booth 7, which enabled Whiley et
al. (1992) to calculate the fruit photosynthesis, and on data of fresh weight
and oil content accumulation of growing ‘Fuerte’ fruits, presented by Ofer
(1986).
The periodical dry matter weight
of fruits, presented in both studies, has been used as “connection numbers”.
That means, that the rate of assimilation, recorded in a ‘Booth 7’ fruit of a
certain dry matter weight, were related to a ‘Fuerte’ fruit with the same dry
matter weight.
The calculations have been made
for fruits during the first period of 10 weeks after anthesis and afterwards
for fruits during 8 periods, of two weeks each, until the 26th week after
anthesis (Table 4).
|
Table 4. Self produced energy of an avocado fruit and its
percentage of the total energy accumulated in a fruit unit. |
|||||
|
(1) Weeks
After Anthesis |
(2) Fruit
Fresh Weight (g) |
(3) Oil in
Fruit (%) |
(4) Fruit Assimilation
of CO2 (nmol·fruit-1·s-1) |
(5) Energy
Produced in 1 Hour Fruit Assimilation (Kcal) |
(6) Fruit
Self assimilation Percentage of Fruit Total Accumulated Energy (%) |
|
10 |
12 |
|
3 |
0.0074 |
21.3 |
|
12 |
32 |
|
10 |
0.0247 |
9.5 |
|
14 |
58 |
2.8 |
18 |
0.0445 |
8.3 |
|
16 |
100 |
2.6 |
30 |
0.0740 |
10.9 |
|
18 |
135 |
3.5 |
38 |
0.0940 |
12.0 |
|
20 |
163 |
4.7 |
47 |
0.1160 |
12.9 |
|
22 |
181 |
7.1 |
50 |
0.1240 |
13.3 |
|
24 |
208 |
7.9 |
53 |
0.1310 |
12.6 |
|
26 |
235 |
9.7 |
55 |
0.1360 |
10.7 |
The figures presented in Table 3,
indicate that there are two main periods during fruit growth, connected to the fruit’s
own photosynthetic activity.
1)
The first ten weeks after anthesis, when the fruit’s
self assimilation provides about 20% of the total energy which accumulates in
the fruit.
2)
The remaining growing period, from the eleventh week
on, when the fruit’s self assimilation provides only about 10% of its
accumulated energy.
This is the place to remind that
in the spring season the assimilation efficiency of most of the previous year
leaves are reduced towards shedding. At that time the new flushing leaves are
still a sink for assimilates rather than a source. During this period a quick
reduction in the starch reservoir level takes place in the avocado tree.
Those facts emphasize the
importance of the self assimilation of the young fruits for their development
and survival.
According to the figures in column
(4) Table 4, the fruit’s self assimilation increases constantly during the
period of fruit growth but its share, out of the total dry matter accumulated
in the fruit, is reduced.
The yearly production and utilization of dry
matter in an avocado tree
Since the monthly assimilation and
the monthly increase in dry matter of an avocado tree have been presented, it
is now possible to calculate the production and utilization of an avocado tree
during a year.
The calculation of the dry matter
production has been made by multiplying the calculated monthly assimilation,
given in mol of CO2, by 30 g (the weight of 1/6 mol glucose) (Table
5).
In order to estimate the
maintenance respiration rate (Rm) of an avocado tree, the weight of an uprooted
‘Hass’ tree, presented by Lovatt (1996), has been used. The utilization of dry
matter for maintenance respiration has been calculated as 3% of the phytomass
of the growing tissues between May and August, 2% during March, April,
September and October, and 1% between November and February. The total monthly utilization
of fry matter of an avocado tree is obtained by adding the monthly Rm to the
monthly growth (Table 6).
|
Table 5. The monthly dry matter (g) production of a
“standard” avocado tree. |
||||||||||||
|
Month: |
Ma |
A |
M |
J |
J |
Au |
S |
O |
N |
D |
J |
F |
|
1. |
1920 |
2445 |
2565 |
3600 |
4872 |
6486 |
6300 |
5280 |
4323 |
3480 |
2835 |
2385 |
|
2. |
|
|
26 |
56 |
172 |
421 |
700 |
791 |
|
|
|
|
|
1+2 |
1920 |
2445 |
2591 |
3656 |
5044 |
6907 |
7000 |
6071 |
4323 |
3480 |
2835 |
2385 |
|
%C |
|
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