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CALCIUM DEFICIENCY SYMPTOMS IN GREENHOUSE CUCUMBERSG. M. WARDResearch Stalion, Agriculture Canada, Harrow, Ontario. Received 28 Mar. j973,accepted 19 lune 1973.

Weno, G. M. 1973. Calcium deficiency symptoms in greenhouse cucumbers.Can. J. Plant Sci. 53: 849-856.In a sand culture experiment with seeded and seedless greenhouse cucumbers(Cucumis sativusL.), induced Ca defciency over a range of nutrient Ca applica-tions produced characteristic foliar and piant symptoms. Severe deficiency wasassociated with top leaf tissue levels of O.20Vo Ca and less. Fruit symptoms were aproportionate stunting, center and stem-end constriction in Burpee hybrid, and aprogressively darker green color in sporu. Mild symptoms were difficult to rec-ognize and could only be confirmed by tissue analysis, although it was not possibleto determine an exact thresbold value for incipient deficiency.

Au cours d'une exp6rience de culture sur sable de concombres de serre avec etsans graines (Cucumis sativusL.),la carence en Ca provoqu6e sur I'ensemble desapplications de Ca s'est traduite par des sympt6mes caract6ristiques sur la feuilleet sur la plante. Aux fortes carences ont correspondu des teneurs de O.2OVoet moins en Ca dans trophie proportionn6e, un 6trang de l,hybiideBurpee, et le vert chez s sympt6mesmineurs qui n'ont pu n,a pas 6t6possible cependant de ce naissante.

INTRODUCTIONIt is well known that the Cucurbitaceae ab-sorb much Ca (Wilkins 1917; Roorda vanEysinga and Smilde 1969; Ward 1967a, b).This element is taken up in larger quantitiesthan any other nutrient, except K and oc-casionally N, bu,t its distribution throughoutthe plant is very uneven. The amounts foundin cucumber (Cucumis sativus L.) fruits arerelatively small. In many plants, lack of Caresults in characteristic deficiency symptoms,which have been described and illustratedand associated with specific Ca concentra-tions in some tissues (Chapman 1966; Wal-lace 1951). However, there is little infor-mation on cucumbers and apparently nodramatic Ca deficiency symptom in the cu-cumber fruit similar to blossomend rot intomatoes (Lycopersicon esculentum Mill.)(Ward 1973) and peppers (Capsicum fru-tescans L.) or corky spots in apples (Malussylvestris Mill.) and some other fruits. Thepresent experiment was designed to deter-mine the effects on cucumber fruits of vary-ing degrees of Ca deficiency and to find whatCa levels are associated with the variousCu. J. Phnt ScL 53: 849-E56 (Oct 1973)

sympto,ms in different tissues, particularlyfruits.MATERIALS AND METIIODSTwo types of cucumber were used in this ex-periment

-the seeded cucumber, cult Burpee

hybrid, and the seedless cucumber, cult Sporu.All tests were made simultaneously with bothtypes. Seedlings were grown from seed in potsof soil and transplanted after 13 days to 4-literpots of soil or washed white sand. Uniformlvsized plants with the first true leaves just show-ing above the cotyledons were selected to pro-vide four replicates per treatment and the rootswere washed free of soil before transplanting.They were watered and fertilized as requiredwith Hewitt's (1966) standard nutrient solutionadjusted to produce the following treatments:meq/liter Ca

Tl: Sand * complete fertilizer(Hewitt 1966)T2: Sand t fertilizer 6T3: Sand * fertilizer 4T4: Sand * fertilizer 2T5: Sand -F fertilizer 0.4T6: Sand t fertilizer without Ca 0T7: Soil f complete fertilizer 8T8: Soil * fertilizer without Ca 0T9: Sand { complete fertilizer for 1 mo,then without Ca.849

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850 CANADIAN JOURNAL OF PLANT SCIENCEThe soil was a light sandy loam used regu-larly in greenhouse culture and it contained 500ppm readily available Ca before fertilization.The plants were grown in a greenhouse underconditions approximating 23 C day temperature,19 C night temperature and 65% relative hu-midity.Ca deficiency symptoms developed on plantsin the various treatments at different times.When they became sufficiently characteristic ofthe disorder, a detailed description was recordedand pictures were taken. Samples were takenfor analysis from relevant treatments, on severaldifferent dates. Selected tissues from two or fourplants were composited, dried in an oven at80 C, and ground. Fruit tissues were preparedon an individual fruit basis and freeze-dried.All samples were analyzed for Ca, K, and Mgby atomic absorption spectrophotometry on adry ash extract. At the end of the growing peri-od the total fruit yields were measured and

recorded.RESULTS AND DISCUSSIONWhereas plants that received an adequatesupply of Ca developed normally andreached a height of 66 cm in 35 days (TableI ), those that were completely deprived ofCa after the 13-day seedling period (T6)quickly developed deflciency symptoms,which in 35 days had the degree of severitydescribed in Table 1, illustrated in Fig. Iand 2, and reflected in the analyses in Table2. These plants continued growing for an-other 6 days but became almost completelynecrotic and were discarded.Plants in T2, T3, and T4 had no visiblefoliar symptoms of deficiency in 35 days buta depression in growth was observed in theform of successive decreases in height andnumber of leaves in T3 and T4. Plants inT5 showed serious deficiency symptoms al-though the growing tips were still viable andexpanding and a few fruit had developed.In 41 days the tops were seriously afiected(Fig. a) and by 49 days had completelyaborted. The plants still remained alive rvitha number of deformed fruit until the ter-mination of the experiment at 67 days.Burpee plants in T2 remained completelynormal in appearance throughout the elperi-ment and the effect of reduced Ca in nutrientapplication was apparent only in the chemi-cal analysis of various tissues. Sporu plantsin T2 began to show mild foliar symptomsof deficiency aftet 49 days (Table 1).

Plants in T3 and T4 demonstrated thatwith decreasing supplies of Ca, deficiencysymptoms appeared sooner and becamemore severe. Decrease in plant height andnumber of leaves was a characteristic fea-ture. Symptoms tended to appear mainly atthe top of the plant unless the deficiency wasextremely severe.Plants in T7 and T8 growing in soil main-tained normal healthy foliage throughout theexperiment, but after 49 days the fruit de-veloping on the upper parts of T8 plants (noadded nutrient Ca) remained small and ab-normal. Plants in T9, which were given ade-quate Ca for I mo after transplanting, de-veloped normally and produced normal fruitduring that period, but when Ca was with-drawn, they responded quickly by develop-ing foliar symptoms near the top of theplant. Any fruit that had set remained smalland deformed and further fruit-set was com-pletely stopped.The analysis of a complete plant of eachcultivar from T6 illustrates the plant re-sponse to complete lack of Ca (Table 2).These figures and those in Table 3 may becompared with previously published datafrom this laboratory (Ward 1967a) showingthe distribution of Ca, K, and Mg in a nor-mal healthy cucumber plant where Ca variedfrom 13.1lVo in the bottom leaf to l.O6Voin the top leaf. Bottom and top leaves fromTl in this experiment were also analyzed(Table 2) and Ca levels are shown to be9.3lVo and l.8lVo for Burpee and' lO.O27oand l.63Vo for Sporu. There was, therefore,a drastic reduction of Ca in all tissues ofT6 plants but the gradient from bottom totop was still maintained. The total quantityof Ca in the whole plant was only 26 mg inBurpee ar,d 2l mg in Sporu. The source ofthis Ca was the seed and the soil in whichthe seedling was grown before being trans-planted to sand.Plants from T5 (Table 3) showed thesame pattern of Ca distribution and gradientas those from T6. They were larger, how-ever, and the Ca levels in the lower leaveswere higher, although percentages in thetop parts of the plant fell to the same lowlevels found in T6. The total quantity ofCa in T5 plants was 46 mg in Burpee and65 mg in Sporu. Nutrient application for

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WARD----CREENHOUSE CUCUMBERS CALCIUM DEFICIENCY 851Table 1. Description of calcium deficiency symptoms on plants of two cucumber cultivars at suc-cessive stases of develooment

Description of plantTreatmentno. Burpee Sporu

T1T2T3T4T5

T6

T7, T8, T9't1, T2

T3T4

T5

T6T7, T8, T9

T1T2T3T4

T5T7T8T9

35 Days .from seed.ingNormal, healthy dark green foliage, Normal, healthy, dark greenht 66 cm, 10 leaves foliage, ht 51 cm, 10 leavesSame as T1 Same as T1Same as T1 except ht 50 cmFoliage normal, ht 31 cm, 7 leavesStunted plants, chlorotic spotting,and interveinal chlorosis on bottomleaves, severe downward curlingon top leaves, small topsPlants severely stunted, shortinternodes, all leaves chlorotic,

top leaves severely curled, topsabortingSame as T1

41 Doys Jrom seedingHt 122 cm, 16 leaves, normalmale and female blossomsSlight leaf curling at topUpper leaves curling,ht 81 cm, 13 leavesThree bottom leaves normal,

remainder chlorotic and curled,small embryo fruit, tops aborting,ht 80 cm, 12 leavesTops necrotic, foliage hard anddying, plants discardedSimilar to T149 Days from seeilingNormal foliage and fruitNormal foliage and fruit

Leaf symptoms on top third of plant,few small fruitSevere plant symptoms,fruit small and deformedSevere symptoms, stunting, abortedtips, fruit small and deformedNormal foliage and fruitNormal foliage, upper fruitssmall and abnormalSymptoms appearing near topof plant, fruit small

Same as T1 except ht 42 cmFoliage normal, ht 25 cm, 8 leavesSame as Burpee

Same as Burpee

Same as T1

Ht 84 cm, 14 leaves, normalmale and female blossomsVery slight leaf curling at topSevere curling of upper leaves andsucker leaves, marginal chlorosis,fruit on suckers,ht 71 cm, 12 leavesSame as Burpee exceptht 70 cm, 11 leavesSame as BurpeeSimilar to T1

Normal foliage and fruitSlight leaf curling and marginalchlorosis near top, normal fruitLeaf symptoms less severe thanBurpee, fruit more normalSevere plant symptoms, plantslightly stunted, fruit slightlyaffectedSevere symptoms, stunting,aborted tips, fruit smallNormal foliage and fruitNormal foliage, some small fruitsSymptoms appearing near top ofplant, fruit deformed and small

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852 cANADIAN JoURNAL oF PLANT scIENcETable 2. Ca, K, and Mg content of some tissues of two cucumber cultivars showing severe Cadeficiency symptoms after 35 days from seeding (treatment T6)

Analysis (7o of dry wt)Burpee Sporu

Description of sample MgagaT6 cotyledonsT6 leaf I (bottom)a

A

67T6 topT6 stem f petioles, bottom halfT6 stem * petioles, top half

T1 leaf 1 (bottom)Tl leaf 9

3.982.341 .591i

.43.25.12.21.12.4.).149.311.81

1.56.93.90J. JJ3.123.242.883.644.254.536. 393.083. 59

.861 .36t.L2I. IJ.91.87.65.91.86.66.491. 78.60

6. 093.20r.42.54.31.11.lo.06.49.15

10.021.63

2.19 l.4r| .77 2 .214.01 r.734.38 1.333.96 . 954.31 .952.88 .672.63 .853.94 .68.J-.)6 .J'3. 16 1.732.88 .45

28 days with a solution containing 8 ppmCa (.4 meq/liter) supplied apProximately44 mg. If the seedlings had the same Cacontent as those in T6, tben the total supplywould amount to 70 mg for Burpee and 65mg for Sporu. These plants, therefore, usedalmost all the available Ca. A normal plantat this stage of growth contains apProxi-mately 500 mg Ca (Ward 1967b).After 4l and 55 days, representative leafsamples from all treatments were analyzed(Table 4). Response to treatment is appar-ent in both bottom and top leaves. Severedeficiency symptoms were associated withtissues with a Ca content of less than .207obut it did not seem to be possible to definea clear threshold value as with Mg (Wardand Miller 1969) or with Ca in tomatoes(Ward 1973) because the symptoms de-veloped gradually and were not strikinglydramatic until they became severe. Calciumdeficiency in cucumber was therefore diffi-cult to diagnose without a chemical analysisof an indicator tissue. The analysis of topleaf tissue of Burpee plants from T9 re-

flected the rapid development of Ca defi-ciency in these plants. Ca was withdrawn43 days after seeding and in 12 days the Cacontent of top leaves had dropped frorn1.54% to .267o.It is apparent that whenthe supply of Ca through the roots was sud-denly cut off, there was no further inputinto the new small leaves at the top. Therewas no mobilization of Ca from other partsof the plant similar to the translocation of Kunder deficiency conditions.After 55 days from seeding, plants in alltreatments had fruits at various stages ofdevelopment and with a variety of deformi-ties. Representative samples that were judgedto be in a condition equivalent to maturitywere picked, photographed (Fig. 5, 6)' de-scribed (Table 5), and analyzed (Table 6).The successive restriction in length andweight with decreasing supplies of Ca fromT1 to T5 is apparent. Decreasing Ca con-centration in the fruit was also found. Theblossom-end striping and stippling seen onthe Burpee fruit in Fig. 5 is a normal geneticcharacteristic, not to be confused with de-

Fig. 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.6. Sporu fruit. T1-T5, 55 days.

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WARD----GREENHOUSE CUCUMBERS CALCIUM DEFICIENCY

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854 CANADIAN JOURNAL OF PLANT SCIENCE

Analysis (/6 of dry wt)Table 3. Ca, K, and Mg content of some tissues of two cucumber cultivars showing severe Cadeficiency symptoms after 4l days from seeding (treatment T5)

Burpee SporuDescription of sample Kaga MgT5 cotyledonsT5 leaf 1 (bottom)

LJA56789

1011l213T5 topT5 small suckersT5 male flowers & embryo fruitT5 stem * petioles, bottom halfT5 stem f petioles, top half

5.804.r73.591.881.26.77.49.23.t4.11.11.12.18.2+.27.41.rl

1. 53. ,J1.99

J.IJ3.623. 553. 70+.76+.LJ3.622.87s.074.523.16+.393.92n 1'

1.031.812.221.621.ZJ.93.o,.49.+7.43.51

. /J.46.ot.41.29

.).504.26r.721.36.93.59.36.25.18.10.09-Lt.ut.11.14.19.19.37.12

| .76 1 .25.61 1.82.86 r.321. 10 r.202.00 t.t41. r0 .862.60 .622,65 .423.77 .512.55 .312.38 .332.95 .462.87 .553.09 .644.50 .733.13 .554.16 .583.98 .424.14 .22flciency symptoms. The center and end con-strictions were caused by Ca deficiency inthis instance, but these expressions are notspecific for Ca alone. They may be pro-duced by other nutrient deficiencies or byfactors that disturb the water balance of thefruit. The seedless cucumber fruit, Sporu,did not show any constriction similar to

Burpee, probably because of the softerthinner skin. However, the Sporu fruit fromT4 and T5 showed a definite darkening inthe green color towards the blossom end.In both cultivars under extreme deficiencythe percentage of Ca reached a very lowlevel in Burpee, .O5Vo, and in Sporu, .06%.In normal fruit (Tl, T7) the Ca levels ap-

Table 4. Ca content of selected sample leaves of two cucumber cultivars af.ter 4l and 55 days fromseedineCa as /s of drY wt

Bottom leaf Top leaf

Treatment Burpee41 days Sporu41 days Burpee41 days Sporu41 days Burpee55 daysT1T2IJT4T5T7T8T9

11.408. 086. 653.952.771r.77tt.8211.38

12.029.418. 08J.A'4.2611.8412.63tt.07

1.731.35.83.58.101.841.581.54

r.74l.J,.88.51.rt2.241.411.61

1.451.48I. JJ.69.114.092.62.26

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WARD----GREENHOUSE CUCUMBERS CALCIUM DEFICIENCYTable 5. Description of calciurn deficiency symptoms on fruit of two cucumber cultivars after 55days from seeding

Description of fruit

855

Treatmentno. Burpee SporuT1T2T3T4T5

Normal green color, slight stripingat blossom end, straightSlightly constricted stem endConstricted center to stem end.lighter color, more stripingCenter severely constricted,very small fruitStunted undeveloped fruit withconstricted blossom end

Light green color, straight orslightly curvedSlightty curved; smaller fruitCurved and shorter than normalCurved fruit, stunted, muchdarker greenSeverely stunted, curved, andvery dark green

proximated those reported in a previousstudy of developing cucumber fruits (Wardand Miller l97O). All fruits in T7 werenormal and only slight abnormalities oc-curred on some fruits in T8. In T9 no fruithad developed on the upper portion of theBurpee plants at the sampling date. TheSporu fruits on the upper parts were de-formed and had a Ca content as low asthose from T5.The interrelationships of K, Ca, and Mgin plant nutrition have been discussed in aprevious paper (Ward 1973). As an addi-tional contribution to this subject the K andMg contents of some tissue samples areshown in Tables 2,3, and 6. The decreasinggradient of Mg from leaf 1 to the top of theplant as reported previously (Ward 1967a)can be seen here. It is also apparent by com-

parison with previous data and by inspeotionof the Tl leaf sample data in Table 2 thatwhen Ca is deficient, the Mg content in-creases. This does not hold for fruit tissues.There is no apparent pattern of relationshipbetween K and Ca.The final assessment of fruit yield after67 days from seeding (Table 7) indicatedthat in this period there was no restrictionin yield of Burpee in the first three treat,ments but a gradual decrease in the yieldof Sporu. The severe effect of Ca deficiencycan be seen in the greatly reduced yieldsfrom T4 and T5. However, ,these plants setas many fruits as those in other treatments,except T5 Sporu, but none of these fruitswere developing. The effect of the diminish-ing supply of Ca in T8 can be seen in thesmall number of additional fruit set prior

Table 6. Ca, K, and Mg content of fruit of two cucumber cultivars frorn all treatments after 55 daysfrom seedingBurpee SporuAnalysis (/6 dty wt) Analysis (0/6 dry wt)Size,Treatment g fresh wt Ca Mgag Size,g fresh wt

T1T2T3T4T5llT8 (lower)T8 (upper)T9

.43 2.55 .18.2r 2.85 . 18.18 2.55 .18.10 2.43 .18.06 2.83 .23

. 54 2.90 .20.40 2.58 .23

.06 2.35 .r7

502184155l16076619619

.45.16.16.20.05.53.42.68

3.582.80s.252.903.402.382.60J.+J

.20.14.18.30.24.21.24.32

52r23r368312109428431264

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856 CANADTAN JoTJRNAL oF pLANT scrENcETable 7. Total yield of fruit from four plants of two cultivars after 67 days from seeding

TotalTreatment no. Totalwt (g) Avg wt Avg length Additional Total(g) (.-) fruit set fruit setl410t6l4t4

I828J0421

1920201510201820

36JJ32282l30JJ30

391263370

2025

7

L720l2421454392

7n84l2

234524242493t926124

3874357234622802885

547381644309

Burpee33530331219362

Sporu484447495255177

I88

100

132l6

88a

1lJ0

131811

T1T2T3T4T5T6T7T8T9

T1T2T3T4T5T6T7T8T9

5089553 12220

t61015l45

to the harvest date, rather than in the yield.And the effect of the sudden stopping of Casupply in T9 is also apparent through theappearance of only one additional fruit inboth cultivars.Characteristic Ca deficiency symptomswere produced in this experiment. The inter-veinal chlorosis and chlorotic spotting, thedownward curling or hooking of leaves, thesevere plant stunting and aborted growingtips are similar to the symptoms found inother species (Wallace 1951; Chapman1966). No specific symptoms were found onfruits other than a proportionate stunting,some constrictions on Burpee, and a pro-gressively darkening green color on Sporu.Extreme deficiency is related to a Ca per-centage in the top leaves below .2OVo, btttno specific percentage can be related to theonset of the disorder.

LITERATURE CITEDCHAPMAN, H. D. 1966. Diagnostic criteriafor plants and soils. Univ. of California Div.Agr. Sci., Riverside, Calif.HEWITT. E. J. 1966. Sand and water culturemethods used in the study of plant nutrition.

2nd ed. Commonwealth Agr. Bur. Tech. Com-mun.22,ROORDA van EYSINGA, J. P. W. L. andSMILDE. K. W. 1969. Nutritional disorders incucumbers and gherkins under glass. Centre forAgricultural Publishing and Documentation,Wageningen, Netherlands. pp. 18-f9.WALLACE, T. 1951. The diagnosis of mineraldeficiencies in plants by visual symptoms. H.M.Stationery Oftice, London.WARD. G. M. 1967a. Greenhouse cucumbernutrition - a growth analysis study. Plant Soil26: 324-332.WARD, G.M. 1967b. Growth and nutrient ab-sorption in greenhouse tomato and cucumber.Proc. Amer. Soc. Hort. Sci. 90: f35-341.WARD, G. M. 1973. Causes of blossom-endrot of tomatoes based on tissue analysis. Can.J. Plant Sci. 53: 169-174WARD, 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. Re-lationship between fruit sizes and nutrient con-tent of greenhouse tomatoes and cucumbers.Can. J. Plant Sci. 50: 451-455.WILKINS, L. K. 1917. The high calcium con-tent of some cucurbit vines. New Jersey Agr.Exp. Sta. Bull. 310: 5-19.

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