growth, physiological processes and yield of tomatoes grown in

PertanikaJ. Trop. Agric. Sci. 18(2): 141-147(1995) ISSN: 0126-6128© Universiti Pertanian Malaysia Press

Growth, Physiological Processes and Yield of Tomatoes Grown in DifferentRoot Zone Volumes Using Sand Culture

MOHD RAZI ISMAIL and S. DALIADepartment of Agronomy and Horticulture,

Universiti Pertanian Malaysia, 43400 Serdang, Selangor, Malaysia

Keywords: tomato, root zone volumes, growth, water relations, mineral nutrition, yield

ABSTRAKPengaruh beberapa isipadu zon akar (2000, 4000 dan 6000 cm3) terhadap pertumbuhan, kaitan air, responsstomata, kadar fotosintesis, pemakanan tanaman dan hasil di dalam kultur pasir. Pokok disampel untukanalisa pertumbuhan setiap 2 minggu selama 56 hari. Perkembangan daun, jisim kering batang dan akardikurangkan dengan isipadu zon akar 2000 cm3 dan 4000 cm3. Hasil basah buah juga dikurangkan apabilatanaman di tanam di dalam isipadu zon akar yang rendah. Walaubagaimanapun, isipadu zon akar tidakmenunjukkan pengaruh yang signiftkan terhadap peratus jisim kering buah dan kandungan bahan terlarut.Pengurangan pertumbuhan dan perkembangan tanaman apabila pokok ditanam didalam isipadu zon akar4000 cm3 dan kurang, dihubungkaitkan dengan pengurangan di dalam potensi air, kadar fotosintesis danpengambilan nutrien.

ABSTRACTThe effects of different root zone volumes (2000, 4000 and 6000 cm3) on growth, water relations, stomatal responses,photosynthetic rate, mineral nutrition and yield of tomatoes (Lycopersicon esculentum Mill.) grown in sandculture were investigated . The plants were sampled for growth analysis fortnightly for a 56-day period. Leaf growth,stem and root dry weights were significanlty reduced with root zone volumes of 2000 and 4000 cm3. Total fruit freshweight was reduced when plants were grown with decreasing root zone volumes. Root zone volumes, however, didnot produce a significant effect on percentage of fruit dry matter and total soluble solids. The reduction in plantgrowth and development was associated with reduction in water potential, photosynthetic rate and mineral nutrientuptake when plants were grown in root zone volumes of 4000 cm3 or less.

INTRODUCTION

Conventional vegetable cultivation in the tropicsis always limited, mainly due to the presence ofsoil-borne diseases and nematodes. Recently,there has been increasing interest in the use ofsoilless substrate technique for producinghorticultural crops, particularly in Malaysia. TheNutrient Film Technique (NFT)-trough systemhas been developed and proven to be feasiblefor producing horticultural crops (Lim, 1990).However, one of the disadvantages of the NFT-trough system still remains, namely therecirculating nutrient solution needs to bemanaged carefully to prevent imbalance of thevarious nutrients. Furthermore, in recirculatingnutrient systems, diseases could easily be spread(Lim, 1986).

To overcome the above problems, researchwas started using a non-recirculating systemutilizing organic and inorganic substrates. Ismailet al. (1993) reported that sand was suitable forthe production of tomatoes, but its disadvantagewas its weight that was very heavy compared withsawdust. Sawdust would be more utilized if therisk of some species of wood producing inhibitoryeffects could be overcome.

One of the main factors that control plantgrowth and development of plants grown usingaggregate soilless culture is the root zone volume.Reducing root zone volume minimises the costof production, especially when utilizing sand.Carmi and Heur (1981) reported that reducingsoil volume was beneficial in producing dwarfplants especially during periods of low radiation.

MOHD RAZI ISMAIL AND S. DALIA

For tomatoes, Raja Harun and Muhamad (1992)found no significant effect of different volumesof coconut potting mix used for the productionof tomatoes. Prasad and Maher (1992) showed asimilar response where varying the volume ofsubstrate from 1.25 Z to 14 / of coarse peat andup to 5 /for rockwool did not significantly affectyield of tomatoes in a recirculating system. Ruffet al (1987) suggested that for a given growingarea, a culture system using small containerswould be more efficient in producing tomatoesof a given weight and size of plant than usinglarge containers based on their data from soilvolumes of 450 cm3 and 13500 cm3. Reducedroot volume which causes root restriction will,however, result in the detrimental effect ofreducing both the morphological andphysiological processes in tomato plants(Peterson et al, 1991).

The present study investigated the possibilityof reducing sand volume for the production oftomatoes with the main objective of increasingproductivity per unit area. The parametersinvestigated were changes in growth, stomatalresponse, photosynthetic rate and mineralnutrition; the possible role of water in affectingplant growth and development; and harvest indexand biomass production were also examined.

MATERIALS AND METHODS

Tomato seeds, (Lycopersicon esculentum Mill. cv.MT1) supplied by the Malaysian AgriculturalResearch and Development Institute (MARDI),were sown in Jiffy pots. Four weeks after sowing,the seedlings were transferred to sand culture.To ensure root establishment, plants wereirrigated with tap water for 4 days. Theexperiment was conducted in a glasshouse atthe Hydroponic Unit, Universiti PertanianMalaysia, Serdang, Selangor, Malaysia. Daily airtemperatures ranged from 24°C to 36°C andrelative humidity between 42% and 84%. A totalof 144 plants was initially grown in three differentsand volumes, namely 2000, 4000 and 6000 cm3,in polybags with holes to allow drainage. Theexperiment was conducted in a completelyrandomized design with six replications. Harvestsof plant material were made on days 14, 28, 42and 56 to estimate net assimilation rate (NAR),harvest index (HI) and root:shoot ratio. Thecalculation of NAR and HI followed those ofHunt (1982). For each harvest, leaf area wasdetermined using an automatic leaf area meter

(Delta T Device, Cambridge, UK). Plants wereseparated into leaves, stems and roots and oven-dried for 72 h at 65°C prior to weighing. Thenutrient concentration of 2.5 mS cm"1 of CooperFormulation (Cooper, 1979) and a pH of 5.5-6.0was supplied to the plants via drip emitters.Fertigation was carried out four times dailyduring the day at intervals of 3 h/per cycle.Each fertigation cycle of 10 min suppliedadequate nutrients and water. Once a week, thesubstrate was flushed with tap water suppliedthrough an emitter to avoid excessiveaccumulation of salts in the substrate. Using thistechnique, we did not observe any toxicitysymptoms throughout the growth period.

Young, fully expanded leaves (3rd to 6th leaffrom top of canopy) were sampled for leaf waterpotential determinations using a pressurechamber (Soil Moisture Eqpt. Corp. U.S.A). Thestomatal resistance was measured with anautomatic porometer (MK3, Delta T Device,Cambridge, UK). Measurement of the netphotosynthetic rate for intact attached leaveswas made using an infra red gas analyser (LGA-2 ADC Hoddesdon, UK).

Nutrient analysis of plant parts was carriedout on plants sampled on days 14 and 42following the procedure outlined by Husni et al(1991). Nitrogen (N), Phosphorus (P), andPotassium (K) contents were determined usingTechnicon Auto-analyser II, while Ca wasdetermined by atomic absorptionspectrophotometer.

Flower number was recorded weekly andpercentage of fruit set determined. Harvestingof fruits was carried out when they changedfrom yellow to red. The numbers of normal andabnormal fruits were recorded. Total fruit weightof normal fruits was obtained. Individual fruitdiameters were recorded using a Vernier caliper.Five fruits per plant were sampled for totalsoluble solids, measured using a handrefractometer, and for percentage dry weight.

RESULTS

Plant Vegetative GrowthOver 56 days, plant height did not differsignificantly (P>0.05) between treatments. Stemdiameter was significantly reduced (P<0.05) inthe 2000 cm3 treatment. No significant differencewas observed for stem diameter between the4000 cm3 and 6000 cm3 treatments (Fig L). Byday 56, leaf area and dry weight were significantly

142 PERTANIKAJ. TROP. AGRIC. SCI. VOL. 18 NO. 2 1995

TOMATOES GROWN IN DIFFERENT ROOT ZONE VOLUMES

Water Relations, Stomatal Resistance (Rs) andPhotosynthetic Rate (Pn)

The leaf water potential declined progressivelywith time. This was particularly marked for thelower root zone volume treatments (Fig. 2), thegreatest drop from -0.29 MPa at 7 days to 0.8MPa at 56 days occurred in the smallest rootzone volume.

60

days after treatments

Fig. 1: Plant height and stem diameter as influenced by differ-ent root zone volumes. 0=2000 cm3 %=4000cw? andA^OOOcm3. Means separatum by DMRT, P=5%

lower (P<0.05) with decreasing volumes of sand(Table 1), being 15% lower in the 2000 cm3

treatment and 30% lower in the 4000 cm3

treatment than for plants grown in 6000 cm3 ofsand. Similar reductions were observed for rootand stem dry weights. The results for root:shootratio between plants grown in diferent root zonevolumes were inconsistent and in general indicatethat there was a shift of dry matter depositionfrom leaves to the stem with decreasing rootzone volume.

Plant Growth Analysis

Table 2 shows root zone volumes had litde or noeffect on net assimilation rate and harvest index.The results for sampling at interval 2-4 weeks forthe net assimilation rate was signifkandy greaterfor the 2000 cms treatment, but this differencebetween treatments diminished with time.

20 30 40 50 60

days after treatments

Fig. 2: Leafivater potential, stomatal resistance and photosyn-

thetic rate as influenced by root zone volumes.

0=200000*; m^OOOcm3 and A=6000cm3. Means

separation by DMRT, P=5%

PERTANIKA J. TROP. AGRIC. SCI. VOL. 18 NO. 2, 1995 143


TABLE 1Influence of root zone volume on leaf area, leaf, root and stem dry weights and root:shoot ratio in MTl

tomato. Data are means of 6 replications for a given days after treatments (DAT). For each growthparameter, means within a row followed by same letter are not significandy different (P>0.05) by DMRT

Parameter Days aftertreatment

2000

Root zone volume (cm3)

4000 6000

Leaf area (cm2)

Leaf dry wt.(g/plant)

Stem dry wt(g/plant)

Root dry wt.(g/plant)

Root:shoot ratio

14284256

149ft-'4256

149ftZo4256

14no

4256

14284256

958140020001796

6.749.69

12.8117.61

3.054.125.136.12

1.311.812.633.04

0.190.190.200.17

cccc

bcbc

bbab

bbbb

aaaa

1286217824682206

8.5712.4315.0621.59

4.095.695.986.11

1.932.433.513.87

0.230.200.230.18

bbbb

ababb

aabaab

ababaab

aaba

1392256229002572

9.5315.2016.7325.34

4.346.046.496.87

2.342.783.404.38

0.240.180.200.17

aaaa

aaaa

aaaa

aaaa

baaa

TABLE 2Net assimilation rate (NAR) and harvest

index (HI) of 'MTl1 tomato plants grownin different root zone volumes. Means

separation by DMRT, P=5%

Parameter

NAR

(g/day/m2)

HI

Root zone

volumes(cm3)

20004000

6000

2000

4000

6000

2-4

11.01

8.84

8.67

0.60

0.61

0.62

a

bb

a

a

a

Interval

4-6

12.70

10.84

9.55

0.76

0.75

0.75

(weeks)

a

a

a

a

a

a

6-8

7.64

7.78

7.26

0.80

0.79

0.83

a

a

a

a

aa

The difference in Rs and Pn between the4000 cm3 and 6000 cm3 treatments was notsignificant throughout the observation period.For the smallest root zone volume treatment, Rswas highest and Pn lowest throughout theobservation period, the Pn at Day 49 being 30%less than for plants grown in 6000 cm3 soil.

Mineral Nutrition

Nutrient content in different plant parts at Day14 and 42 for the three treatments are shown inTable 3. N content in young leaves on bothsampling dates was significantly lower in plantsgrown in 2000 cm3 sand. In contrast, the differentroot zone volumes had no appreciable effect onN content in the roots and fruits.

Reducing root zone volume to 4000 cm3

did not significantly affect P content in youngleaves sampled at Day 14, but caused aremarkable decline of P content by Day 42.



TABLE 3Nutrient content (g/plant) of different parts of tomato plants grown for 14 and 42 days in different rootzone volumes. For each nutrient, means within a column followed by the same letter are not significantly

different (P>0.05) by DMRT

Nutrient

N

P

K

Ca

Root zone

volume(cm3)

200040006000

2000

40006000

2000

4000

6000

2000

4000

6000

Youngleaves

0.16b0.27a0.28a

0.05b0.07a

0.07a

0.23b

0.21ab

0.33a

0.06b

0.09a

0.11a

14 days

Roots

0.02a0.02a0.03a

0.01b

0.01b

0.03a

0.01a

0.01a

0.02a

0.01a

0.02a

0.02a

Youngleaves

0.46b0.63a0.76a

0.17c0.19b

0.26a

0.58b

0.65ab

0.68a

0.27b

0.37a

0.41a

42

Matureleaves

0,41c0.52b0.60a

0.19b

0.20b

0.23a

0.51b

0.60a

0.61a

0.38b

0.42b

0.58a

days

Roots

0.06b0.06b0.07a

0.04b0.04b

0.06a

0.05ab0.04b

0.06a

0.04a

0.04a

0.04a

Fruits

0.78a0.80a0.88a

0.29b

0.33b0.41a

1.12b

1.28ab

1.60a

0.21b0.23ab

0.30a

However, differences in P content between the2000 cm3 and 4000 cm3 treatments were notsignificant for root, mature leaves and fruit. Asimilar reduction in K content was found inyoung leaves of plants grown in the smallestroot zone volume. There was also a 40%reduction in Ca content in young leaves in the2000 cm3 treatment compared with plants grownin 6000 cm3 sand; but for Ca content in theroot, there was a negligible difference.

Yield

Table 4 shows the effects of the different rootzone volumes on fruit production at final harvest.The total fruit fresh weight was 30% and 18%greater in the 6000 cm3 treatment than in the2000 cm3 and 4000 cm3 treatments, respectively.Reduction in fruit fresh weight in the 2000 cm3

treatment was associated with lower fruit numbercaused by reduction in percentage fruit set, andsmaller diameter fruits. Harvest index measuredon a dry weight basis was not significandy differentbetween treatments (Table 2), nor did root zonevolumes affect percentage dry matter and totalsoluble solids in the fruits.

DISCUSSIONReducing root zone volume caused an alterationin the pattern of growth and development oftomato plants grown in sand culture. The studyindicated that plants grown at the lowest rootzone volume suffered stress in leaf photosyntheticrate through partial or complete stomatal closure,which could have been brought about by thereduction in leaf internal water relations asindicated by lowered leaf water potential withreduced root zone volumes. For alder seedlings,Tschaplinski and Blake (1985) also demonstratedthat reduction in internal leaf water potentialcaused stomatal closure in plants grown underrestricted root conditions. Decline in leaf waterpotential and increases in stomatal resistancealso correlate with termination of leaf growth(Shulze, 1986) In contrast, for soybean plants,Krizek et al (1985) reported that restricted rootzone volume did not induce water stress andthat reduction in leaf growth was associated withother physiological mechanisms. In the presentstudy, although reduction in leaf water potentialfor the lowest root zone volume did not exceed-1.0 MPa, we suggest that this level may have

PERTANIKAJ. TROP. AGRIC. SCI. VOL. 18 NO, 2, 1995 145


TABLE 4Influence of different root zone volumes on fruit production. Means separation by DMRT, 5%

Fruit number (unit/plant)

% fruit set

% normal fruit

Fruit diameter (cm)

Fruit fresh wt (g/plant)

% fruit dry matter

Total soluble solids (% Brix)

Days aftertreatments

-

-

_

28

42

56

Total

-

-

2000

15.0 b

68.3 b

85.1 a

3.43 b

84.72 b

398.15 a

407.24 b

811.11 c

8.42 a

5.67 a

Root zone volume

4000

19.0 a

84.0 a

85.7 a

3.55 ab

152.25 a

435.92 a

454.98 b

1043.15 b

7.92 a

6.33 a

(cm3)

6000

19.0 a

86.5 a

84.3 a

3.83 a

156.30 a

494.19 a

616.04 a

1266.53 a7.12 a5.67 a

already affected leaf growth. This is consistentwith the suggestion that the primary effects ofwater stress (slight to moderate) are reducedleaf growth either in the cell extention phase orin both the cell division and cell extention phasesof leaf growth (Begg and Turner, 1976).

The other mechanisms brought into play byrestricted root growth include a combination ofreduced water absorption, reduced hormonesynthesis and an increased allocation ofassimilates translocated to the root. The lattermechanism may be applicable in the presentsudy, where rootrshoot ratio tended to be greaterwith reduction in root zone volume from 6000to 4000 cm3. The involvement of gibberellin andcytokinin has been demonstrated by Richardsand Rowe (1977) and Carmi et al (1983).Peterson et al (1991) demonstrated a slightincrease in ethylene production whenadventitious rooting was initiated, but that overallethylene production rates did not differsignificantly in tomato plants. Similarly, Thomas(1993) ruled out the only possible regulation ofethylene production that caused the reductionin plant size and alterations in allometry ofcarrots plants, as being due to both nutrientdepletion and lack of aeration or restrictedrecirculated nutrient solution. No attempt wasmade in the present study to examine the roleof hormonal changes due to reduced root zonevolume.

The fact that reduced root zone volumedid not decrease the harvest index agrees withthat observed by Ruff et al (1987), and indicatesthat tomato plants can adjust their fruit productionper plant according to pressures on the rootsystem, a similar situation to those grown in highplanting densities. Furthermore, the shift of drymatter allocation to the root and probably also tothe stem, contributes toward this effect and mayexplain the lack of significant difference in thenet assimilation ratio.

The lower concentration of nutrient elementsin the plant with reducing root zone volumes mayreflect a physiological disruption that limits nutrientuptake, but we have no data to support this.

Reducing root volume can be a practicalmeans of reducing cost of production of tomatoesusing soilless media. However, this study showsthat yield was reduced by 34% at the lowest rootzone volume (2000 cm3). Further studies need tobe carried out to minimize or eliminate thereduction in yield by adjusting aspects of irrigationand nutrition requirements.

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(Received 18 December 1993; accepted 1 July 1995)

PERTANIKAJ. TROP. AGRIC. SCI. VOL. 18 NO. 2, 1995 147

growth, physiological processes and yield of tomatoes grown in

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