Manganese Toxicity in Avocado (Persea
Jonathan
Edward Tracy
Candidate for degree of
Master of Science in Soil Science from
SUMMARY
Avocado trees exposed to increasing concentrations of
solution Mn suffered from increased leaf abscission. Tissue concentrations of
Mn, K, and P were increased, while Fe decreased in response to solution Mn.
Approximately 1.1% of the avocado groves surveyed
produced leaf samples with Mn concentrations greater than 1,000 mg/kg. Death of
young, field grown avocado trees is associated with Mn concentrations of
2,000-2,500 mg/kg in six- to ten-month old leaves, and Mn in the saturation
extract of 0.8-4.4 mg/l. Surviving trees were found to have Mn concentrations
of 0.03-0.3 mg/l in the saturation extract. Toxic concentrations of soil Mn
were associated with heavy fertilization-acidification of a manganiferous
soil, or with the addition of a wood-product soil amendment high in soluble Mn.
Approximately 6.8% of the avocado groves surveyed
produced leaf samples with P concentrations below 0.8 g/kg on a dry weight
basis, which is less than optimum.
Introduction
Many of the virgin soils utilized for avocado
production in
Mn has been observed to accumulate in avocado leaf
tissue throughout the growing season under acid soil conditions, but stable
leaf Mn values have been observed under more favorable soil conditions. Data
from salt tolerance studies have demonstrated a wide variation in the ability
of various avocado varieties to accumulate Mn.
Soil conditions favoring Mn solubility (and plant
availability) are low pH and Eh (reducing conditions). Sulfate solubilizes more Mn than either
chloride or nitrate, and Mn is displaced from cation
exchange sites by cations in the order Mg > Ca > NH4=Na >
K.
Materials and Methods
Forty-eight avocado seedlings (Persea
The nutrient solutions were pumped continuously for
15 minutes four times daily onto the top of the plastic buckets. The nutrient
solution contained the following concentrations of ions expressed in mg/l: 113
NO3-N, 33 P04-P, 100 K, 100 Ca, 30 Mg, 67 SO4-S,
5.0 Fe (chelate), 0.1 B, 0.1 Mn, 0.1 Zn, 0.05 Cu,
0.01 Mo. The SO4-S concentration increased throughout the experiment
due to adjustment of the solutions to pH 4.0 ± 0.5 with H2SO4.
Six weeks after transplanting, the nutrient solutions
were amended with MnSO4-4H2O to yield a metal
concentration of either 0.0, 2.0, 4.0, or 10.0 mg/l in
a specified tank. Thirteen weeks after transplanting, the nutrient solutions
were renewed; and at sixteen weeks, the Mn concentrations were doubled to 0.0,
4.0, 8.0, or 20.0 mg/l.
Twenty-eight weeks after transplanting, the trees
were harvested and divided into top leaves, bottom leaves, stems, and roots;
and dry weights were determined (other growth parameters are reported
elsewhere).
The top portions of the trees were analyzed for K,
Ca, Mg, Zn, Fe, Mn, Cu, and Al by atomic absorption spectrophotometry;
for N, P, and B by colorimetry; and for S by
turbidity.
Field Data
Manganese was determined on 1,444 avocado tissue
samples from commercial groves in northern
Two additional groves were analyzed in detail by the
above methods with N by microkjeldahl and soil
analysis by CFA methods.
Results and Discussion
Sand Culture Experiment:
Reduced measurements of leaves and roots were
obtained in response to Mn in solution, although only leaf area and dry weight of
bottom leaves were significantly reduced at the 0.05 level of probability. The
relationship between total dry weight and Mn in solution is presented in Fig.
1. The increased statistical error at higher Mn concentrations is the result of
individual trees largely defoliating, while stronger individuals in the same
treatments continued to maintain most of their leaves. Leaves suffering from Mn
toxicity were not observed to be smaller than comparable leaves from untreated
trees.
The appearance of Mn
toxicity symptoms on the lower surface of avocado leaves is shown in Fig. 2,
along with the concentration of Mn in the respective
nutrient solutions. The blackening of the blade surface immediately adjacent to
the midrib and larger veins is evident. Occasionally, the symptoms are
expressed on only part of an affected leaf; but the black color in relation to
the veins has been characteristic in every case observed to date. This symptom
is easily confused with normal leaf senescence in which the identical tissue
turns dark brown. Average chemical composition of the bottom leaves from trees
grown with Mn-amended nutrient solutions are
presented in Table 1.
At harvest, an internal blackening of cambium tissue
was noted in the trunks of some of the trees exposed to the higher
concentrations of Mn.

Fig. 1. Total plant dry weight as a function of solution Mn.
Survey Data
The distribution of Mn in 1,444 samples from
commercial avocado groves in northern

Fig. 2. The appearance of Mn toxicity symptoms on the bottom
surface of avocado (eaves as a
/unction of solution Mn
concentration.
|
Table
1. Average composition of six to
ten month old avocado leaves with respect to solution Mn. |
||||||
|
Solution Mn,
mg/L |
g/kg |
|||||
|
N |
P |
K |
Ca |
Mg |
S |
|
|
0.1 |
19.9 |
1.3 |
10.1 |
18.5 |
4.6 |
2.2 |
|
4 |
18.5 |
1.3 |
10.1 |
18.0 |
4.0 |
2.1 |
|
8 |
20.7 |
1.5 |
12.1 |
17.6 |
3.9 |
2.3 |
|
20 |
20.3 |
1.7 |
12.8 |
16.5 |
3.8 |
2.5 |
|
|
mg/kg |
|||||
|
|
Zn |
Fe |
Mn |
Cu |
B |
Al |
|
0.1 |
110 |
146 |
297 |
8.3 |
68 |
67 |
|
4 |
83 |
130 |
2025 |
6.7 |
58 |
77 |
|
8 |
90 |
123 |
3700 |
6.3 |
54 |
65 |
|
20 |
95 |
99 |
6300 |
7.7 |
60 |
51 |

Fig. 3. Frequency
distribution of Mn concentrations (dry wt. basis) in leaf samples of commercial
avocado groves in
Mn Toxicity in Poorly-drained Mn-rich Soils
This particular grove is situated at the apex of a
steep hill in
The analytical data presented in Table 2 represent two
separate samplings of the grove.
The April sampling consisted of parallel samples
collected from the north and south sides of the hill. Leaves were selected on
the basis of exhibiting toxicity symptoms from the healthiest looking trees.
The concentrations of Mn, Al, and Mg are elevated in these leaves, while B and
K are depressed. Symptoms of B or K deficiency were not observed.
The leaves from the May sampling were confined to the
south slope and consisted of mature, normal looking leaves, except where
younger leaves were the only choice available (95% defoliation). This has
resulted in analyses which have been confounded by the effect of leaf age in
addition to the effects of Mn. These analyses are still useful in that they
suggest that K and B may be in adequate supply to the tree, even though they
are depressed in the older leaves.
|
Table 2. Leaf analysis of Mn-affected
avocado trees grown on a shallow, rocky, heavy-textured and poorly drained
soil (Fallbrook Ag-Lab Inc., ICP data courtesy Dr. G. Bradford). |
|||||||||||||
|
|
g/kg |
|
mg/kg |
||||||||||
|
Sample* |
N |
P |
K |
Ca |
Mg |
Na |
|
Zn |
Fe |
Mn |
Cu |
B |
Mn/Fe |
|
NS 4/81 |
20 |
1.5 |
4.1 |
20 |
7.0 |
|
|
24 |
110 |
2000 |
2.5 |
20 |
18.2 |
|
SS 4/81 |
20 |
1.4 |
5.4 |
19 |
6.0 |
|
|
24 |
. 90 |
2500 |
2.5 |
20 |
27.8 |
|
Not D 5/81 |
27 |
2.5 |
12.0 |
10 |
3.8 |
0.3 |
|
26 |
55 |
1450 |
2.0 |
30 |
26.4 |
|
50% D 5/81 |
35 |
3.8 |
17.0 |
12 |
5.0 |
0.9 |
|
49 |
87 |
1150 |
2.0 |
130 |
13.2 |
|
95% D 5/81 |
40 |
4.0 |
17.0 |
11 |
4.8 |
1.5 |
|
48 |
110 |
530 |
2.5 |
60 |
4.8 |
|
|
mg/kg |
|
|
|
Less than 5 mg/kg |
||||||||
|
|
Al |
Si |
Sr |
Ba |
|
|
|
Ag, Be, Cd, Co, Cr, Ga, Li,
Mo, Ni, Pb, Sn, Ti |
|||||
|
NS 4/81 ICP |
250 |
138 |
14 |
77 |
|
|
|
||||||
|
*. NS is the north slope,
SS is the south slope, D is defoliated and ICP is Inductively coupled plasma
spectroscopy. |
|||||||||||||
N in the stem is indicated as having a nearly
significant positive correlation with solution Mn (Table 2). Whether this
represents N which has simply been displaced from the roots by Mn (i.e.,
facilitating translocation), or whether this is an active accumulation of N in
response to excessive Mn, is not clear.
Nonsignificant negative correlations were obtained for Ca and Mg
with Mn in solution. A significant negative correlation was obtained with Ca
but not Mg in coffee trees (Pavan, 1979), and
decreases in both Ca and Mg concentration with increasing Mn are discernible in
the apple leaf data of Eggert and Hayden (1970). Mengel and Kirkby (1978) suggest
that Mn-Ca-Mg effects are the result of competing ions being restricted by the
overall ionic balance of the plant.
The mortality rate in this grove was approximately
80% of the avocado trees, two years after planting.
Mn Toxicity Related to the use of Soil Amendments
This one year old grove is situated on a gentle,
south facing slope in
Mn toxicity symptoms were visible in this grove and
were confirmed by leaf analysis (Table 3). Soil analysis showed a close
correlation between soil pH and tree condition. Large amounts of water soluble
Mn were present in a wood product soil amendment applied to the grove at the
time of planting (Table 3).
The mortality rate in this grove was approximately
25% of the avocado trees one year after planting.
|
Table 3. Leaf and soil analysis of a Mn
affected avocado grove (courtesy Fallbrook Ag-Lab Inc.) |
|||||||||||||
|
|
P |
K |
Na |
Cl |
|
Zn |
Fe |
Mn |
Cu |
Mn1 |
|
Mne |
pH |
|
Sample |
g/kg |
|
mg/kg |
|
mg/L |
||||||||
|
Top Leaves |
|
|
|
|
|
|
|
|
|
|
|
|
|
|
Healthy
trees |
3.8 |
19.2 |
0.2 |
2.0 |
|
37 |
70 |
232 |
3 |
|
|
|
|
|
Weak
trees |
3.2 |
24.0 |
0.4 |
5.0 |
|
49 |
97 |
785 |
3 |
|
|
|
|
|
Bottom Leaves |
|
|
|
|
|
|
|
|
|
|
|
|
|
|
Healthy
trees |
1.2 |
6.4 |
0.2 |
4.0 |
|
17 |
85 |
950 |
2 |
|
|
|
|
|
Weak
trees |
1.3 |
3.9 |
0.1 |
7.0 |
|
16 |
100 |
2285 |
1 |
|
|
|
|
|
Soil Samples |
|
|
|
|
|
|
|
|
|
|
|
|
|
|
Native
soil |
|
|
|
|
|
|
|
|
|
182 |
|
0.03 |
6.2 |
|
Healthy
trees |
|
|
|
|
|
|
|
|
|
|
|
|
6.4 |
|
Weak
trees |
|
|
|
|
|
|
|
|
|
|
|
|
4.9 |
|
Wood
product soil amendment |
|
|
|
|
|
|
|
|
|
180 |
|
1.2 |
5.7 |
|
1 Determined
on a H2O2-H2SO4 digest. |
|||||||||||||
Literature Cited
Bohn, H. L., B. L. McNeal and G. A. O'Conner. 1979. Soil Chemistry. John Wiley and Sons, Inc., pp
210-211.
Bowman, R. H. 1973. Soil Survey of the San
Gustafson, C. D., A. W. Marsh, R. L. Branson and
Hamilton, W. R., A. R. Woolley
and A. C. Bishop. 1974. A guide
to Minerals, Rocks and Fossils. Crescent Books,
Rodgers,
T. H. 1965. Geologic Map of