Calcium Cucumber 3r5319
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CALCIUM DEFICIENCY SYMPTOMS IN GREENHOUSE CUCUMBERS G. M. WARD Research Stalion, Agriculture Canada, Harrow, Ontario. Received 28 Mar. j973, accepted 19 lune 1973.
Weno, G.
M. 1973. Calcium deficiency
symptoms
Can. J. Plant Sci. ed from pubs.aic.ca by 94.201.235.211 on 04/28/13 For personal use only.
Can. J. Plant Sci. 53: 849-856.
in
greenhouse cucumbers.
In a sand culture experiment with seeded and seedless greenhouse cucumbers (Cucumis sativusL.), induced Ca defciency over a range of nutrient Ca applications produced characteristic foliar and piant symptoms. Severe deficiency was associated with top leaf tissue levels of O.20Vo Ca and less. Fruit symptoms were a proportionate stunting, center and stem-end constriction in Burpee hybrid, and a progressively darker green color in sporu. Mild symptoms were difficult to recognize and could only be confirmed by tissue analysis, although it was not possible to determine an exact thresbold value for incipient deficiency. Au cours d'une exp6rience de culture sur sable de concombres de serre avec et
sans graines (Cucumis sativusL.),la carence en Ca provoqu6e sur I'ensemble des applications de Ca s'est traduite par des sympt6mes caract6ristiques sur la feuille et sur la plante. Aux fortes carences ont correspondu des teneurs de O.2OVo
et moins en Ca dans portionn6e, un 6trang Burpee, et le vert chez mineurs qui n'ont pu possible cependant
trophie pro
de l,hybiide s sympt6mes n,a pas 6t6 ce naissante.
de
INTRODUCTION
sympto,ms
It is well
known that the Cucurbitaceae absorb much Ca (Wilkins 1917; Roorda van Eysinga and Smilde 1969; Ward 1967a, b). This element is taken up in larger quantities than any other nutrient, except K and occasionally N, bu,t its distribution throughout the plant is very uneven. The amounts found in cucumber (Cucumis sativus L.) fruits are relatively small. In many plants, lack of Ca results in characteristic deficiency symptoms,
which have been described and illustrated and associated with specific Ca concentra-
tions in some tissues (Chapman 1966; Wal-
lace 1951). However, there is little information on cucumbers and apparently no dramatic Ca deficiency symptom in the cu-
fruit similar to blossomend rot in tomatoes (Lycopersicon esculentum Mill.)
in different tissues, particularly
fruits.
MATERIALS AND METIIODS Two types of cucumber were used in this experiment the seeded cucumber, cult Burpee
- the seedless cucumber, cult Sporu. hybrid, and All
tests were made simultaneously with both types. Seedlings were grown from seed in pots of soil and transplanted after 13 days to 4-liter pots of soil or washed white sand. Uniformlv sized plants with the first true leaves just showing above the cotyledons were selected to provide four replicates per treatment and the roots were washed free of soil before transplanting. They were watered and fertilized as required with Hewitt's (1966) standard nutrient solution adjusted to produce the following treatments:
meq/liter Ca
cumber
Tl:
Sand
(Ward 1973) and peppers (Capsicum frutescans L.) or corky spots in apples (Malus sylvestris Mill.) and some other fruits. The
T2: T3:
Sand t fertilizer Sand * fertilizer Sand * fertilizer Sand -F fertilizer Sand t fertilizer without Ca Soil f complete fertilizer
present experiment was designed to determine the effects on cucumber fruits of varying degrees of Ca deficiency and to find what
Ca levels are associated with the various Cu. J. Phnt
ScL 53: 849-E56
(Oct
*
complete fertilizer
(Hewitt 1966)
T4: T5: T6: T7: T8: Soil * fertilizer without Ca T9: Sand { complete fertilizer for then without Ca.
1973)
849
6
4 2
0.4 0 8
0 1 mo,
850
CANADIAN JOURNAL OF PLANT SCIENCE
The soil was a light sandy loam used regularly in greenhouse culture and it contained 500 ppm readily available Ca before fertilization. The plants were grown in a greenhouse under conditions approximating 23 C day temperature, 19 C night temperature and 65% relative humidity. Ca deficiency symptoms developed on plants in the various treatments at different times. When they became sufficiently characteristic of
Can. J. Plant Sci. ed from pubs.aic.ca by 94.201.235.211 on 04/28/13 For personal use only.
the disorder, a detailed description was recorded and pictures were taken. Samples were taken for analysis from relevant treatments, on several
different dates. Selected tissues from two or four plants were composited, dried in an oven at 80 C, and ground. Fruit tissues were prepared on an individual fruit basis and freeze-dried. All samples were analyzed for Ca, K, and Mg by atomic absorption spectrophotometry on a dry ash extract. At the end of the growing period the total fruit yields were measured and recorded.
RESULTS AND DISCUSSION Whereas plants that received an adequate
supply of Ca developed normally and reached a height of 66 cm in 35 days (Table those that were completely deprived of Ca after the 13-day seedling period (T6) I
),
quickly developed deflciency
symptoms,
which in 35 days had the degree of severity described in Table 1, illustrated in Fig. I
and 2, and reflected in the analyses in Table 2. These plants continued growing for an-
other 6 days but became almost completely necrotic and were discarded. Plants in T2, T3, and T4 had no visible foliar symptoms of deficiency in 35 days but a depression in growth was observed in the
form of successive decreases in height and number of leaves in T3 and T4. Plants in T5 showed serious deficiency symptoms although the growing tips were still viable and expanding and a few fruit had developed. In 41 days the tops were seriously afiected (Fig. a) and by 49 days had completely aborted. The plants still remained alive rvith a number of deformed fruit until the termination of the experiment at 67 days. Burpee plants in T2 remained completely normal in appearance throughout the elperiment and the effect of reduced Ca in nutrient application was apparent only in the chemical analysis of various tissues. Sporu plants in T2 began to show mild foliar symptoms of deficiency aftet 49 days (Table 1).
Plants in T3 and T4 demonstrated that with decreasing supplies of Ca, deficiency symptoms appeared sooner and became more severe. Decrease in plant height and number of leaves was a characteristic feature. Symptoms tended to appear mainly at the top of the plant unless the deficiency was extremely severe. Plants in T7 and T8 growing in soil maintained normal healthy foliage throughout the experiment, but after 49 days the fruit developing on the upper parts of T8 plants (no added nutrient Ca) remained small and abnormal. Plants in T9, which were given adequate Ca for I mo after transplanting, developed normally and produced normal fruit during that period, but when Ca was withdrawn, they responded quickly by developing foliar symptoms near the top of the plant. Any fruit that had set remained small and deformed and further fruit-set was completely stopped. The analysis of a complete plant of each
cultivar from T6 illustrates the plant response to complete lack of Ca (Table 2). These figures and those in Table 3 may be
compared with previously published data from this laboratory (Ward 1967a) showing the distribution of Ca, K, and Mg in a normal healthy cucumber plant where Ca varied from 13.1lVo in the bottom leaf to l.O6Vo in the top leaf. Bottom and top leaves from
Tl in this experiment were also analyzed (Table 2) and Ca levels are shown to be
9.3lVo and l.8lVo for Burpee and' lO.O27o and l.63Vo for Sporu. There was, therefore, a drastic reduction of Ca in all tissues of T6 plants but the gradient from bottom to top was still maintained. The total quantity
of Ca in the whole plant was only 26 mg in Burpee ar,d 2l mg in Sporu. The source of this Ca was the seed and the soil in which the seedling was grown before being transplanted to sand. Plants from T5 (Table 3) showed the
of Ca distribution and gradient as those from T6. They were larger, however, and the Ca levels in the lower leaves were higher, although percentages in the same pattern
top parts of the plant fell to the same low levels found in T6. The total quantity of Ca in T5 plants was 46 mg in Burpee and 65 mg in Sporu. Nutrient application for
WARD----CREENHOUSE CUCUMBERS CALCIUM DEFICIENCY
Table
1. Description of calcium deficiency
851
symptoms on plants of two cucumber cultivars at suc-
cessive stases of develooment
Description of plant Treatment no.
Burpee
Sporu
35 Days .from seed.ing
T1
Normal, healthy dark green ht 66 cm, 10 leaves
T2
Same as T1 Same as T1 except
Can. J. Plant Sci. ed from pubs.aic.ca by 94.201.235.211 on 04/28/13 For personal use only.
T3
T4 T5
T6
ht
foliage,
50 cm
Foliage normal, ht 31 cm, 7 leaves Stunted plants, chlorotic spotting, and interveinal chlorosis on bottom leaves, severe downward curling on top leaves, small tops Plants severely stunted, short internodes, all leaves chlorotic, top leaves severely curled, tops
Normal, healthy, dark green foliage, ht 51 cm, 10 leaves Same as T1 Same as T1 except
ht 42 cm Foliage normal, ht 25 cm, 8 leaves Same as Burpee
Same as Burpee
aborting T7, T8, T9
Same as T1
't1, T2
Ht
T3 T4
Same as T1
41 Doys Jrom seeding 122 cm, 16 leaves, normal male and female blossoms
Slight leaf curling at top Upper leaves curling,
ht
81 cm, 13 leaves
T5
Three bottom leaves normal, remainder chlorotic and curled, small embryo fruit, tops aborting,
T6
Tops necrotic, foliage hard and dying, plants discarded Similar to T1
ht
T7, T8, T9
Ht
84 cm, 14 leaves, normal male and female blossoms Very slight leaf curling at top Severe curling of upper leaves and sucker leaves, marginal chlorosis, fruit on suckers,
ht 71 cm, 12 leaves Same as Burpee except ht 70 cm, 11 leaves
80 cm, 12 leaves Same as Burpee
Similar to T1
49 Days from seeiling
T1 T2
Normal foliage and fruit Normal foliage and fruit
T3
Leaf symptoms on top third of plant,
Normal foliage and fruit Slight leaf curling and marginal chlorosis near top, normal fruit Leaf symptoms less severe than
T4
Severe plant symptoms,
Severe plant symptoms, plant
few small fruit
fruit small and deformed T5 T7 T8 T9
Severe symptoms, stunting, aborted
tips, fruit small and deformed Normal foliage and fruit Normal foliage, upper fruits
small and abnormal Symptoms appearing near top of plant, fruit small
Burpee, fruit more normal
slightly stunted, fruit slightly affected Severe symptoms, stunting,
aborted tips, fruit small Normal foliage and fruit Normal foliage, some small fruits Symptoms appearing near top of plant, fruit deformed and small
852 Table
cANADIAN JoURNAL
oF PLANT scIENcE
cucumber cultivars showing severe Ca deficiency symptoms after 35 days from seeding (treatment T6)
2. Ca, K, and Mg content of some tissues of two
Analysis (7o of dry wt) Sporu
Burpee
Description of sample
T6 cotyledons T6 leaf I (bottom)
Can. J. Plant Sci. ed from pubs.aic.ca by 94.201.235.211 on 04/28/13 For personal use only.
a
6 7
Tl
K
3.98 2.34
1.56 .93
1
.59 1i .43
3.12
.25
3.24 2.88 3.64 4.25 4.53
1
A
T6 T6 T6 T1
Ca
.12 .21 .12 .4.)
top stem f petioles, bottom half stem * petioles, top half leaf 1 (bottom)
.14 9.31
leaf 9
1.81
Mg
Ca
.86 .36
6. 09
.90
t.L2
r.42
J. JJ
I. IJ .91
.54
.87 .65
3.20 .31 .11
.lo
.91
.06
6. 39
.86 .66 .49
.15
3.08
1. 78
10.02
3. 59
.60
1.63
28 days with a solution containing 8 ppm Ca (.4 meq/liter) supplied apProximately 44 mg. If the seedlings had the same Ca content as those in T6, tben the total supply would amount to 70 mg for Burpee and 65 mg for Sporu. These plants, therefore, used almost all the available Ca. A normal plant at this stage of growth contains apProximately 500 mg Ca (Ward 1967b). After 4l and 55 days, representative leaf samples from all treatments were analyzed (Table 4). Response to treatment is apparent in both bottom and top leaves. Severe deficiency symptoms were associated with tissues with a Ca content of less than .207o but it did not seem to be possible to define a clear threshold value as with Mg (Ward and Miller 1969) or with Ca in tomatoes (Ward 1973) because the symptoms developed gradually and were not strikingly dramatic until they became severe. Calcium deficiency in cucumber was therefore difficult to diagnose without a chemical analysis of an indicator tissue. The analysis of top leaf tissue of Burpee plants from T9 re-
.49
K
2.19 | .77 4.01 4.38 3.96 4.31 2.88 2.63 3.94 .J-.)6 3.
16 2.88
Mg
l.4r 2
.21
r.73 1.33 .
95
.95 .67 .85 .68 .J'
1.73 .45
flected the rapid development of Ca deficiency in these plants. Ca was withdrawn 43 days after seeding and in 12 days the Ca content of top leaves had dropped frorn
1.54% to .267o.It is apparent that when the supply of Ca through the roots was sud-
denly cut off, there was no further input into the new small leaves at the top. There was no mobilization of Ca from other parts
of the plant similar to the translocation of K under deficiency conditions.
After 55 days from seeding, plants in all treatments had fruits at various stages of development and with a variety of deformities. Representative samples that were judged to be in a condition equivalent to maturity were picked, photographed (Fig. 5, 6)' described (Table 5), and analyzed (Table 6). The successive restriction in length and weight with decreasing supplies of Ca from T1 to T5 is apparent. Decreasing Ca concentration in the fruit was also found. The blossom-end striping and stippling seen on the Burpee fruit in Fig. 5 is a normal genetic characteristic, not to be confused with de-
1. Severe Ca deficiency with necrotic growing tip. T6, 35 days. Fig. 2. Normal plant Tl (pesticide spray residue on leaves) and T6 plant 35 days. Fig. 3. Normal plants T1' 4l days. Flg. 4. Seveie Ca deficiency. T5, 4l days. Fig. 5. Burpee hybrid fruit. T1-T5, 55 days. Fig. Fig.
6. Sporu
fruit.
T1-T5, 55 days.
Can. J. Plant Sci. ed from pubs.aic.ca by 94.201.235.211 on 04/28/13 For personal use only.
WARD----GREENHOUSE CUCUMBERS CALCIUM DEFICIENCY
CANADIAN JOURNAL OF PLANT SCIENCE
854
Table
3. Ca, K, and Mg content of some tissues of two cucumber cultivars showing deficiency symptoms after 4l days from seeding (treatment T5)
severe Ca
Analysis (/6 of dry wt) Sporu
Burpee
Description of sample
T5 cotyledons T5 leaf 1 (bottom)
Can. J. Plant Sci. ed from pubs.aic.ca by 94.201.235.211 on 04/28/13 For personal use only.
5.80
1. 53
1.03
.).50
4.r7
. ,J
1.81
4.26
1.99
2.22 1.62
1.36
7
.23
+.76
.49
.36 .25
8
.t4
+.LJ
9
.11 .11 .12
3.62 2.87
.+7 .43
.18 .10
.51
.09 -Lt
A
5
10 11
J.IJ
3.62 3. 55 3. 70
s.07
.93
.o,
.ut
13
.11
.18 .2+ .27
. /J
3.16
.46
.14 .19
.ot
.19
n 1'
.rl
flciency symptoms. The center and end constrictions were caused by Ca deficiency in this instance, but these expressions are not specific for Ca alone. They may be produced by other nutrient deficiencies or by factors that disturb the water balance of the
fruit. The seedless cucumber fruit, Sporu, did not show any constriction similar to 4.
4.52 +.39 3.92
.41
.76 .61 .86 1. 10 2.00 1. r0 2.60 2,65 3.77 2.55 2.38 2.95 2.87 3.09 4.50 3.13 4.16 3.98 4.14
.59
l2 top small suckers male flowers & embryo fruit stem * petioles, bottom half stem f petioles, top half
Table
1.ZJ .93
.41
.37
.29
.12
Mg
|
r.72
6
J
K
Ca
3.59 1.88 1.26 .77 .49
L
T5 T5 T5 T5 T5
Mg
Ca
Burpee, probably because
1
.25
1.82
r.32 r.20
t.t4 .86 .62 .42 .51 .31
.33 .46 .55
.64 .73
.55 .58 .42 .22
of the softer
thinner skin. However, the Sporu fruit from T4 and T5 showed a definite darkening in the green color towards the blossom end. In both cultivars under extreme deficiency the percentage of Ca reached a very low level in Burpee, .O5Vo, and in Sporu, .06%. In normal fruit (Tl, T7) the Ca levels ap-
Ca content of selected sample leaves of two cucumber cultivars af.ter
4l and 55 days
from
seedine
Ca as /s of drY wt
Top leaf
Bottom leaf Treatment T1 T2 IJ
T4 T5 T7 T8 T9
Sporu
Burpee 41 days
Sporu
Burpee 55 days
Burpee 41 days
41 days
11.40
12.02
8. 08 6. 65
9.41
1.73 1.35
8. 08
.83
.88
I. JJ
3.95 2.77
J.A'
.51
.69
1r.77
11.84
.58 .10 1.84 1.58 1.54
tt.82
11.38
4.26 12.63
tt.07
41 days
r.74 l.J,
1.45 1.48
.rt
.11
2.24
4.09
1.41 1.61
2.62 .26
855
WARD----GREENHOUSE CUCUMBERS CALCIUM DEFICIENCY
5. Description of calciurn
Table
deficiency symptoms on fruit of two cucumber cultivars after 55 days from seeding Description of fruit
Treatment no.
T1 T2 T3
Can. J. Plant Sci. ed from pubs.aic.ca by 94.201.235.211 on 04/28/13 For personal use only.
Sporu
Burpee
T4 T5
Normal green color, slight striping at blossom end, straight Slightly constricted stem end Constricted center to stem end. lighter color, more striping
Light green color, straight or
Center severely constricted, very small fruit Stunted undeveloped fruit with constricted blossom end
Curved fruit, stunted, much darker green Severely stunted, curved, and
proximated those reported
in a previous
and Miller l97O). All fruits in T7 were normal and only slight abnormalities occurred on some fruits in T8. In T9 no fruit had developed on the upper portion of the Burpee plants at the sampling date. The
when Ca is deficient, the Mg content increases. This does not hold for fruit tissues. There is no apparent pattern of relationship between
as
those from T5.
The interrelationships of K, Ca, and Mg in plant nutrition have been discussed in a previous paper (Ward 1973). As an additional contribution to this subject the K and
Mg contents of some tissue samples are shown in Tables 2,3, and 6. The decreasing
except T5 Sporu, but none of these fruits were developing. The effect of the diminishing supply of Ca in T8 can be seen in the
gradient of Mg from leaf 1 to the top of the plant as reported previously (Ward 1967a) can be seen here. It is also apparent by com-
6.
Ca, K, and Mg content of
K and Ca.
The final assessment of fruit yield after 67 days from seeding (Table 7) indicated that in this period there was no restriction in yield of Burpee in the first three treat, ments but a gradual decrease in the yield of Sporu. The severe effect of Ca deficiency can be seen in the greatly reduced yields from T4 and T5. However, ,these plants set as many fruits as those in other treatments,
Sporu fruits on the upper parts were de-
Table
very dark green parison with previous data and by inspeotion of the Tl leaf sample data in Table 2 that
study of developing cucumber fruits (Ward
formed and had a Ca content as low
slightly curved Slightty curved; smaller fruit Curved and shorter than normal
small number
of
additional
fruit set prior
fruit of two cucumber cultivars frorn all treatments after
55 days
from seeding Burpee
Sporu
Analysis (/6 dty wt)
Treatment
Size,
g fresh wt
Ca
T1 T2 T3
502 184 155
T4
l1
T5
60 766 196
.45 .16 .16 .20 .05
19
ll
T8 (lower) T8 (upper) T9
Analysis (0/6 dry wt) Size,
Mg
g fresh wt
.20 .14 .18 .30 .24
368 312 109
.53 .42
3.58 2.80 s.25 2.90 3.40 2.38 2.60
.21
428
.24
431
.68
J.+J
.32
52r
23r
264
Ca
.43 .2r .18 .10 .06 . 54 .40 .06
K
2.55 2.85 2.55 2.43 2.83 2.90 2.58 2.35
Mg .18 . 18
.18 .18 .23 .20 .23
.r7
856
CANADTAN JoTJRNAL
Table
oF pLANT
scrENcE
7. Total yield of fruit from four plants of two cultivars
Treatment
Total no.
Total
wt
wt (g)
Avg
(g)
after 67 days from seeding
length Additional (.-) fruit set
Avg
Total
fruit
set
Can. J. Plant Sci. ed from pubs.aic.ca by 94.201.235.211 on 04/28/13 For personal use only.
Burpee
T1
I
T2 T3 T4 T5 T6 T7 T8 T9
8
2345 2424 2493
335 303
19
312
20
t926
193
124
62
15 10
13
5089
2l
553
391 263 370
8 10
7 n
l4
8 4
t6
l2
t4
20
10
l4
0 1
6
2220
T1 T2
8 8
T3 T4 T5 T6 T7 T8 T9
a
1l
3874 3572 3462 2802
J
20 25
20 18
Sporu 484
20
I
7
36
8
t6
447
JJ
2
10
495 255
32
8
15
28
J
l4
885
177
2l
0
5
13 18
5473 8164
421
30
4
L7
454
JJ
2
20
11
4309
392
30
1
l2
0
to the harvest date, rather than in the yield. And the effect of the sudden stopping of Ca supply in T9 is also apparent through the appearance of only one additional fruit in both cultivars. Characteristic Ca deficiency symptoms were produced in this experiment. The interveinal chlorosis and chlorotic spotting, the downward curling or hooking of leaves, the severe plant stunting and aborted growing tips are similar to the symptoms found in
other species (Wallace 1951;
Chapman
1966). No specific symptoms were found on
fruits other than a proportionate stunting, some constrictions on Burpee, and a progressively darkening green color on Sporu. Extreme deficiency is related to a Ca percentage in the top leaves below .2OVo, bttt no specific percentage can be related to the onset of the disorder. LITERATURE CITED CHAPMAN, H. D. 1966. Diagnostic criteria for plants and soils. Univ. of California Div. Agr. Sci., Riverside, Calif.
HEWITT. E. J. 1966. Sand and water culture methods used in the study of plant nutrition.
2nd ed. Commonwealth Agr. Bur. Tech. Commun.22,
ROORDA van EYSINGA, J. P. W. L. and SMILDE. K. W. 1969. Nutritional disorders in cucumbers and gherkins under glass. Centre for Agricultural Publishing and Documentation, Wageningen, Netherlands. pp. 18-f9. WALLACE, T. 1951. The diagnosis of mineral deficiencies in plants by visual symptoms. H.M. Stationery Oftice, London.
WARD. G.
M.
1967a. Greenhouse cucumber
nutrition a growth analysis study. Plant Soil 26: 324-332. WARD, G.M. 1967b. Growth and nutrient absorption in greenhouse tomato and cucumber. Proc. Amer. Soc. Hort. Sci. 90: f35-341.
M. 1973. Causes of blossom-end tomatoes based on tissue analysis. Can. J. Plant Sci. 53: 169-174
WARD, G.
rot of
WARD, G. M. and MILLER, M.
J.
1969.
Magnesium deficiency in greenhouse tomatoes. Can. J. Plant Sci. 49: 5f-59. WARD, G. M. and MILLER, M. J. 1970. Relationship between fruit sizes and nutrient content of greenhouse tomatoes and cucumbers. Can. J. Plant Sci. 50: 451-455. WILKINS, L. K. 1917. The high calcium content of some cucurbit vines. New Jersey Agr. Exp. Sta. Bull. 310: 5-19.
Weno, G.
M. 1973. Calcium deficiency
symptoms
Can. J. Plant Sci. ed from pubs.aic.ca by 94.201.235.211 on 04/28/13 For personal use only.
Can. J. Plant Sci. 53: 849-856.
in
greenhouse cucumbers.
In a sand culture experiment with seeded and seedless greenhouse cucumbers (Cucumis sativusL.), induced Ca defciency over a range of nutrient Ca applications produced characteristic foliar and piant symptoms. Severe deficiency was associated with top leaf tissue levels of O.20Vo Ca and less. Fruit symptoms were a proportionate stunting, center and stem-end constriction in Burpee hybrid, and a progressively darker green color in sporu. Mild symptoms were difficult to recognize and could only be confirmed by tissue analysis, although it was not possible to determine an exact thresbold value for incipient deficiency. Au cours d'une exp6rience de culture sur sable de concombres de serre avec et
sans graines (Cucumis sativusL.),la carence en Ca provoqu6e sur I'ensemble des applications de Ca s'est traduite par des sympt6mes caract6ristiques sur la feuille et sur la plante. Aux fortes carences ont correspondu des teneurs de O.2OVo
et moins en Ca dans portionn6e, un 6trang Burpee, et le vert chez mineurs qui n'ont pu possible cependant
trophie pro
de l,hybiide s sympt6mes n,a pas 6t6 ce naissante.
de
INTRODUCTION
sympto,ms
It is well
known that the Cucurbitaceae absorb much Ca (Wilkins 1917; Roorda van Eysinga and Smilde 1969; Ward 1967a, b). This element is taken up in larger quantities than any other nutrient, except K and occasionally N, bu,t its distribution throughout the plant is very uneven. The amounts found in cucumber (Cucumis sativus L.) fruits are relatively small. In many plants, lack of Ca results in characteristic deficiency symptoms,
which have been described and illustrated and associated with specific Ca concentra-
tions in some tissues (Chapman 1966; Wal-
lace 1951). However, there is little information on cucumbers and apparently no dramatic Ca deficiency symptom in the cu-
fruit similar to blossomend rot in tomatoes (Lycopersicon esculentum Mill.)
in different tissues, particularly
fruits.
MATERIALS AND METIIODS Two types of cucumber were used in this experiment the seeded cucumber, cult Burpee
- the seedless cucumber, cult Sporu. hybrid, and All
tests were made simultaneously with both types. Seedlings were grown from seed in pots of soil and transplanted after 13 days to 4-liter pots of soil or washed white sand. Uniformlv sized plants with the first true leaves just showing above the cotyledons were selected to provide four replicates per treatment and the roots were washed free of soil before transplanting. They were watered and fertilized as required with Hewitt's (1966) standard nutrient solution adjusted to produce the following treatments:
meq/liter Ca
cumber
Tl:
Sand
(Ward 1973) and peppers (Capsicum frutescans L.) or corky spots in apples (Malus sylvestris Mill.) and some other fruits. The
T2: T3:
Sand t fertilizer Sand * fertilizer Sand * fertilizer Sand -F fertilizer Sand t fertilizer without Ca Soil f complete fertilizer
present experiment was designed to determine the effects on cucumber fruits of varying degrees of Ca deficiency and to find what
Ca levels are associated with the various Cu. J. Phnt
ScL 53: 849-E56
(Oct
*
complete fertilizer
(Hewitt 1966)
T4: T5: T6: T7: T8: Soil * fertilizer without Ca T9: Sand { complete fertilizer for then without Ca.
1973)
849
6
4 2
0.4 0 8
0 1 mo,
850
CANADIAN JOURNAL OF PLANT SCIENCE
The soil was a light sandy loam used regularly in greenhouse culture and it contained 500 ppm readily available Ca before fertilization. The plants were grown in a greenhouse under conditions approximating 23 C day temperature, 19 C night temperature and 65% relative humidity. Ca deficiency symptoms developed on plants in the various treatments at different times. When they became sufficiently characteristic of
Can. J. Plant Sci. ed from pubs.aic.ca by 94.201.235.211 on 04/28/13 For personal use only.
the disorder, a detailed description was recorded and pictures were taken. Samples were taken for analysis from relevant treatments, on several
different dates. Selected tissues from two or four plants were composited, dried in an oven at 80 C, and ground. Fruit tissues were prepared on an individual fruit basis and freeze-dried. All samples were analyzed for Ca, K, and Mg by atomic absorption spectrophotometry on a dry ash extract. At the end of the growing period the total fruit yields were measured and recorded.
RESULTS AND DISCUSSION Whereas plants that received an adequate
supply of Ca developed normally and reached a height of 66 cm in 35 days (Table those that were completely deprived of Ca after the 13-day seedling period (T6) I
),
quickly developed deflciency
symptoms,
which in 35 days had the degree of severity described in Table 1, illustrated in Fig. I
and 2, and reflected in the analyses in Table 2. These plants continued growing for an-
other 6 days but became almost completely necrotic and were discarded. Plants in T2, T3, and T4 had no visible foliar symptoms of deficiency in 35 days but a depression in growth was observed in the
form of successive decreases in height and number of leaves in T3 and T4. Plants in T5 showed serious deficiency symptoms although the growing tips were still viable and expanding and a few fruit had developed. In 41 days the tops were seriously afiected (Fig. a) and by 49 days had completely aborted. The plants still remained alive rvith a number of deformed fruit until the termination of the experiment at 67 days. Burpee plants in T2 remained completely normal in appearance throughout the elperiment and the effect of reduced Ca in nutrient application was apparent only in the chemical analysis of various tissues. Sporu plants in T2 began to show mild foliar symptoms of deficiency aftet 49 days (Table 1).
Plants in T3 and T4 demonstrated that with decreasing supplies of Ca, deficiency symptoms appeared sooner and became more severe. Decrease in plant height and number of leaves was a characteristic feature. Symptoms tended to appear mainly at the top of the plant unless the deficiency was extremely severe. Plants in T7 and T8 growing in soil maintained normal healthy foliage throughout the experiment, but after 49 days the fruit developing on the upper parts of T8 plants (no added nutrient Ca) remained small and abnormal. Plants in T9, which were given adequate Ca for I mo after transplanting, developed normally and produced normal fruit during that period, but when Ca was withdrawn, they responded quickly by developing foliar symptoms near the top of the plant. Any fruit that had set remained small and deformed and further fruit-set was completely stopped. The analysis of a complete plant of each
cultivar from T6 illustrates the plant response to complete lack of Ca (Table 2). These figures and those in Table 3 may be
compared with previously published data from this laboratory (Ward 1967a) showing the distribution of Ca, K, and Mg in a normal healthy cucumber plant where Ca varied from 13.1lVo in the bottom leaf to l.O6Vo in the top leaf. Bottom and top leaves from
Tl in this experiment were also analyzed (Table 2) and Ca levels are shown to be
9.3lVo and l.8lVo for Burpee and' lO.O27o and l.63Vo for Sporu. There was, therefore, a drastic reduction of Ca in all tissues of T6 plants but the gradient from bottom to top was still maintained. The total quantity
of Ca in the whole plant was only 26 mg in Burpee ar,d 2l mg in Sporu. The source of this Ca was the seed and the soil in which the seedling was grown before being transplanted to sand. Plants from T5 (Table 3) showed the
of Ca distribution and gradient as those from T6. They were larger, however, and the Ca levels in the lower leaves were higher, although percentages in the same pattern
top parts of the plant fell to the same low levels found in T6. The total quantity of Ca in T5 plants was 46 mg in Burpee and 65 mg in Sporu. Nutrient application for
WARD----CREENHOUSE CUCUMBERS CALCIUM DEFICIENCY
Table
1. Description of calcium deficiency
851
symptoms on plants of two cucumber cultivars at suc-
cessive stases of develooment
Description of plant Treatment no.
Burpee
Sporu
35 Days .from seed.ing
T1
Normal, healthy dark green ht 66 cm, 10 leaves
T2
Same as T1 Same as T1 except
Can. J. Plant Sci. ed from pubs.aic.ca by 94.201.235.211 on 04/28/13 For personal use only.
T3
T4 T5
T6
ht
foliage,
50 cm
Foliage normal, ht 31 cm, 7 leaves Stunted plants, chlorotic spotting, and interveinal chlorosis on bottom leaves, severe downward curling on top leaves, small tops Plants severely stunted, short internodes, all leaves chlorotic, top leaves severely curled, tops
Normal, healthy, dark green foliage, ht 51 cm, 10 leaves Same as T1 Same as T1 except
ht 42 cm Foliage normal, ht 25 cm, 8 leaves Same as Burpee
Same as Burpee
aborting T7, T8, T9
Same as T1
't1, T2
Ht
T3 T4
Same as T1
41 Doys Jrom seeding 122 cm, 16 leaves, normal male and female blossoms
Slight leaf curling at top Upper leaves curling,
ht
81 cm, 13 leaves
T5
Three bottom leaves normal, remainder chlorotic and curled, small embryo fruit, tops aborting,
T6
Tops necrotic, foliage hard and dying, plants discarded Similar to T1
ht
T7, T8, T9
Ht
84 cm, 14 leaves, normal male and female blossoms Very slight leaf curling at top Severe curling of upper leaves and sucker leaves, marginal chlorosis, fruit on suckers,
ht 71 cm, 12 leaves Same as Burpee except ht 70 cm, 11 leaves
80 cm, 12 leaves Same as Burpee
Similar to T1
49 Days from seeiling
T1 T2
Normal foliage and fruit Normal foliage and fruit
T3
Leaf symptoms on top third of plant,
Normal foliage and fruit Slight leaf curling and marginal chlorosis near top, normal fruit Leaf symptoms less severe than
T4
Severe plant symptoms,
Severe plant symptoms, plant
few small fruit
fruit small and deformed T5 T7 T8 T9
Severe symptoms, stunting, aborted
tips, fruit small and deformed Normal foliage and fruit Normal foliage, upper fruits
small and abnormal Symptoms appearing near top of plant, fruit small
Burpee, fruit more normal
slightly stunted, fruit slightly affected Severe symptoms, stunting,
aborted tips, fruit small Normal foliage and fruit Normal foliage, some small fruits Symptoms appearing near top of plant, fruit deformed and small
852 Table
cANADIAN JoURNAL
oF PLANT scIENcE
cucumber cultivars showing severe Ca deficiency symptoms after 35 days from seeding (treatment T6)
2. Ca, K, and Mg content of some tissues of two
Analysis (7o of dry wt) Sporu
Burpee
Description of sample
T6 cotyledons T6 leaf I (bottom)
Can. J. Plant Sci. ed from pubs.aic.ca by 94.201.235.211 on 04/28/13 For personal use only.
a
6 7
Tl
K
3.98 2.34
1.56 .93
1
.59 1i .43
3.12
.25
3.24 2.88 3.64 4.25 4.53
1
A
T6 T6 T6 T1
Ca
.12 .21 .12 .4.)
top stem f petioles, bottom half stem * petioles, top half leaf 1 (bottom)
.14 9.31
leaf 9
1.81
Mg
Ca
.86 .36
6. 09
.90
t.L2
r.42
J. JJ
I. IJ .91
.54
.87 .65
3.20 .31 .11
.lo
.91
.06
6. 39
.86 .66 .49
.15
3.08
1. 78
10.02
3. 59
.60
1.63
28 days with a solution containing 8 ppm Ca (.4 meq/liter) supplied apProximately 44 mg. If the seedlings had the same Ca content as those in T6, tben the total supply would amount to 70 mg for Burpee and 65 mg for Sporu. These plants, therefore, used almost all the available Ca. A normal plant at this stage of growth contains apProximately 500 mg Ca (Ward 1967b). After 4l and 55 days, representative leaf samples from all treatments were analyzed (Table 4). Response to treatment is apparent in both bottom and top leaves. Severe deficiency symptoms were associated with tissues with a Ca content of less than .207o but it did not seem to be possible to define a clear threshold value as with Mg (Ward and Miller 1969) or with Ca in tomatoes (Ward 1973) because the symptoms developed gradually and were not strikingly dramatic until they became severe. Calcium deficiency in cucumber was therefore difficult to diagnose without a chemical analysis of an indicator tissue. The analysis of top leaf tissue of Burpee plants from T9 re-
.49
K
2.19 | .77 4.01 4.38 3.96 4.31 2.88 2.63 3.94 .J-.)6 3.
16 2.88
Mg
l.4r 2
.21
r.73 1.33 .
95
.95 .67 .85 .68 .J'
1.73 .45
flected the rapid development of Ca deficiency in these plants. Ca was withdrawn 43 days after seeding and in 12 days the Ca content of top leaves had dropped frorn
1.54% to .267o.It is apparent that when the supply of Ca through the roots was sud-
denly cut off, there was no further input into the new small leaves at the top. There was no mobilization of Ca from other parts
of the plant similar to the translocation of K under deficiency conditions.
After 55 days from seeding, plants in all treatments had fruits at various stages of development and with a variety of deformities. Representative samples that were judged to be in a condition equivalent to maturity were picked, photographed (Fig. 5, 6)' described (Table 5), and analyzed (Table 6). The successive restriction in length and weight with decreasing supplies of Ca from T1 to T5 is apparent. Decreasing Ca concentration in the fruit was also found. The blossom-end striping and stippling seen on the Burpee fruit in Fig. 5 is a normal genetic characteristic, not to be confused with de-
1. Severe Ca deficiency with necrotic growing tip. T6, 35 days. Fig. 2. Normal plant Tl (pesticide spray residue on leaves) and T6 plant 35 days. Fig. 3. Normal plants T1' 4l days. Flg. 4. Seveie Ca deficiency. T5, 4l days. Fig. 5. Burpee hybrid fruit. T1-T5, 55 days. Fig. Fig.
6. Sporu
fruit.
T1-T5, 55 days.
Can. J. Plant Sci. ed from pubs.aic.ca by 94.201.235.211 on 04/28/13 For personal use only.
WARD----GREENHOUSE CUCUMBERS CALCIUM DEFICIENCY
CANADIAN JOURNAL OF PLANT SCIENCE
854
Table
3. Ca, K, and Mg content of some tissues of two cucumber cultivars showing deficiency symptoms after 4l days from seeding (treatment T5)
severe Ca
Analysis (/6 of dry wt) Sporu
Burpee
Description of sample
T5 cotyledons T5 leaf 1 (bottom)
Can. J. Plant Sci. ed from pubs.aic.ca by 94.201.235.211 on 04/28/13 For personal use only.
5.80
1. 53
1.03
.).50
4.r7
. ,J
1.81
4.26
1.99
2.22 1.62
1.36
7
.23
+.76
.49
.36 .25
8
.t4
+.LJ
9
.11 .11 .12
3.62 2.87
.+7 .43
.18 .10
.51
.09 -Lt
A
5
10 11
J.IJ
3.62 3. 55 3. 70
s.07
.93
.o,
.ut
13
.11
.18 .2+ .27
. /J
3.16
.46
.14 .19
.ot
.19
n 1'
.rl
flciency symptoms. The center and end constrictions were caused by Ca deficiency in this instance, but these expressions are not specific for Ca alone. They may be produced by other nutrient deficiencies or by factors that disturb the water balance of the
fruit. The seedless cucumber fruit, Sporu, did not show any constriction similar to 4.
4.52 +.39 3.92
.41
.76 .61 .86 1. 10 2.00 1. r0 2.60 2,65 3.77 2.55 2.38 2.95 2.87 3.09 4.50 3.13 4.16 3.98 4.14
.59
l2 top small suckers male flowers & embryo fruit stem * petioles, bottom half stem f petioles, top half
Table
1.ZJ .93
.41
.37
.29
.12
Mg
|
r.72
6
J
K
Ca
3.59 1.88 1.26 .77 .49
L
T5 T5 T5 T5 T5
Mg
Ca
Burpee, probably because
1
.25
1.82
r.32 r.20
t.t4 .86 .62 .42 .51 .31
.33 .46 .55
.64 .73
.55 .58 .42 .22
of the softer
thinner skin. However, the Sporu fruit from T4 and T5 showed a definite darkening in the green color towards the blossom end. In both cultivars under extreme deficiency the percentage of Ca reached a very low level in Burpee, .O5Vo, and in Sporu, .06%. In normal fruit (Tl, T7) the Ca levels ap-
Ca content of selected sample leaves of two cucumber cultivars af.ter
4l and 55 days
from
seedine
Ca as /s of drY wt
Top leaf
Bottom leaf Treatment T1 T2 IJ
T4 T5 T7 T8 T9
Sporu
Burpee 41 days
Sporu
Burpee 55 days
Burpee 41 days
41 days
11.40
12.02
8. 08 6. 65
9.41
1.73 1.35
8. 08
.83
.88
I. JJ
3.95 2.77
J.A'
.51
.69
1r.77
11.84
.58 .10 1.84 1.58 1.54
tt.82
11.38
4.26 12.63
tt.07
41 days
r.74 l.J,
1.45 1.48
.rt
.11
2.24
4.09
1.41 1.61
2.62 .26
855
WARD----GREENHOUSE CUCUMBERS CALCIUM DEFICIENCY
5. Description of calciurn
Table
deficiency symptoms on fruit of two cucumber cultivars after 55 days from seeding Description of fruit
Treatment no.
T1 T2 T3
Can. J. Plant Sci. ed from pubs.aic.ca by 94.201.235.211 on 04/28/13 For personal use only.
Sporu
Burpee
T4 T5
Normal green color, slight striping at blossom end, straight Slightly constricted stem end Constricted center to stem end. lighter color, more striping
Light green color, straight or
Center severely constricted, very small fruit Stunted undeveloped fruit with constricted blossom end
Curved fruit, stunted, much darker green Severely stunted, curved, and
proximated those reported
in a previous
and Miller l97O). All fruits in T7 were normal and only slight abnormalities occurred on some fruits in T8. In T9 no fruit had developed on the upper portion of the Burpee plants at the sampling date. The
when Ca is deficient, the Mg content increases. This does not hold for fruit tissues. There is no apparent pattern of relationship between
as
those from T5.
The interrelationships of K, Ca, and Mg in plant nutrition have been discussed in a previous paper (Ward 1973). As an additional contribution to this subject the K and
Mg contents of some tissue samples are shown in Tables 2,3, and 6. The decreasing
except T5 Sporu, but none of these fruits were developing. The effect of the diminishing supply of Ca in T8 can be seen in the
gradient of Mg from leaf 1 to the top of the plant as reported previously (Ward 1967a) can be seen here. It is also apparent by com-
6.
Ca, K, and Mg content of
K and Ca.
The final assessment of fruit yield after 67 days from seeding (Table 7) indicated that in this period there was no restriction in yield of Burpee in the first three treat, ments but a gradual decrease in the yield of Sporu. The severe effect of Ca deficiency can be seen in the greatly reduced yields from T4 and T5. However, ,these plants set as many fruits as those in other treatments,
Sporu fruits on the upper parts were de-
Table
very dark green parison with previous data and by inspeotion of the Tl leaf sample data in Table 2 that
study of developing cucumber fruits (Ward
formed and had a Ca content as low
slightly curved Slightty curved; smaller fruit Curved and shorter than normal
small number
of
additional
fruit set prior
fruit of two cucumber cultivars frorn all treatments after
55 days
from seeding Burpee
Sporu
Analysis (/6 dty wt)
Treatment
Size,
g fresh wt
Ca
T1 T2 T3
502 184 155
T4
l1
T5
60 766 196
.45 .16 .16 .20 .05
19
ll
T8 (lower) T8 (upper) T9
Analysis (0/6 dry wt) Size,
Mg
g fresh wt
.20 .14 .18 .30 .24
368 312 109
.53 .42
3.58 2.80 s.25 2.90 3.40 2.38 2.60
.21
428
.24
431
.68
J.+J
.32
52r
23r
264
Ca
.43 .2r .18 .10 .06 . 54 .40 .06
K
2.55 2.85 2.55 2.43 2.83 2.90 2.58 2.35
Mg .18 . 18
.18 .18 .23 .20 .23
.r7
856
CANADTAN JoTJRNAL
Table
oF pLANT
scrENcE
7. Total yield of fruit from four plants of two cultivars
Treatment
Total no.
Total
wt
wt (g)
Avg
(g)
after 67 days from seeding
length Additional (.-) fruit set
Avg
Total
fruit
set
Can. J. Plant Sci. ed from pubs.aic.ca by 94.201.235.211 on 04/28/13 For personal use only.
Burpee
T1
I
T2 T3 T4 T5 T6 T7 T8 T9
8
2345 2424 2493
335 303
19
312
20
t926
193
124
62
15 10
13
5089
2l
553
391 263 370
8 10
7 n
l4
8 4
t6
l2
t4
20
10
l4
0 1
6
2220
T1 T2
8 8
T3 T4 T5 T6 T7 T8 T9
a
1l
3874 3572 3462 2802
J
20 25
20 18
Sporu 484
20
I
7
36
8
t6
447
JJ
2
10
495 255
32
8
15
28
J
l4
885
177
2l
0
5
13 18
5473 8164
421
30
4
L7
454
JJ
2
20
11
4309
392
30
1
l2
0
to the harvest date, rather than in the yield. And the effect of the sudden stopping of Ca supply in T9 is also apparent through the appearance of only one additional fruit in both cultivars. Characteristic Ca deficiency symptoms were produced in this experiment. The interveinal chlorosis and chlorotic spotting, the downward curling or hooking of leaves, the severe plant stunting and aborted growing tips are similar to the symptoms found in
other species (Wallace 1951;
Chapman
1966). No specific symptoms were found on
fruits other than a proportionate stunting, some constrictions on Burpee, and a progressively darkening green color on Sporu. Extreme deficiency is related to a Ca percentage in the top leaves below .2OVo, bttt no specific percentage can be related to the onset of the disorder. LITERATURE CITED CHAPMAN, H. D. 1966. Diagnostic criteria for plants and soils. Univ. of California Div. Agr. Sci., Riverside, Calif.
HEWITT. E. J. 1966. Sand and water culture methods used in the study of plant nutrition.
2nd ed. Commonwealth Agr. Bur. Tech. Commun.22,
ROORDA van EYSINGA, J. P. W. L. and SMILDE. K. W. 1969. Nutritional disorders in cucumbers and gherkins under glass. Centre for Agricultural Publishing and Documentation, Wageningen, Netherlands. pp. 18-f9. WALLACE, T. 1951. The diagnosis of mineral deficiencies in plants by visual symptoms. H.M. Stationery Oftice, London.
WARD. G.
M.
1967a. Greenhouse cucumber
nutrition a growth analysis study. Plant Soil 26: 324-332. WARD, G.M. 1967b. Growth and nutrient absorption in greenhouse tomato and cucumber. Proc. Amer. Soc. Hort. Sci. 90: f35-341.
M. 1973. Causes of blossom-end tomatoes based on tissue analysis. Can. J. Plant Sci. 53: 169-174
WARD, G.
rot of
WARD, G. M. and MILLER, M.
J.
1969.
Magnesium deficiency in greenhouse tomatoes. Can. J. Plant Sci. 49: 5f-59. WARD, G. M. and MILLER, M. J. 1970. Relationship between fruit sizes and nutrient content of greenhouse tomatoes and cucumbers. Can. J. Plant Sci. 50: 451-455. WILKINS, L. K. 1917. The high calcium content of some cucurbit vines. New Jersey Agr. Exp. Sta. Bull. 310: 5-19.
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