Copyright is owned by the Author of the thesis. Permission is given for a copy to be downloaded by an individual for the purpose of research and private study only. The thesis may not be reproduced elsewhere without the permission of the Author. Title of thesis: Massey University Library Thesis Copyright Form (1) (a) I give permission for my thesis to be made available to readers in Massey University Library under conditions determined by the Librarian. (b) I do not wish my thesis to be made available to readers without my written consent for . . . months. ' (2) (a) I agree that my thesis, or a copy, may be sent to another institution under conditions determined by the Librarian. (b) I do not wish my thesis, or a copy, to be sent to another institution without my written consent for months. (3) (a) I agree that my thesis may be copied for Library use. (b) I do not wish my thesis to be copied for Library use for ... months. Signed The copyright of this thesis belongs to the author. Readers must sign their name in the space below to show that they recognise this. They are asked to add their permanent address. NAME AND ADDRESS SJ~C{,o,o -pa~ \\Cl0 f-CJ~\:J\11)11 577n'e, Co~ od/\ o~ V\ \tu 1/V\ ti wy') is f \rvi Gf?,vws ·1o50 ·DATE THE EFFECTS OF HIGH CONDUCTIVITY LIQUID FEEDS ON THE YIELD AND QUALITY OF OUTDOOR GROWN TOMATOES A Thesis Presented in Partial Fulfilment of the Requirements for the Degree of Master of Horticultural Science in Vegetable Production at Massey University Palmerston North, New Zealand EDGARDO F. FADALlAN Department of Plant Science 1993 ABSTRACT Studies were conducted to evaluate the effects of high conductivity liquid feeds applied using drip irrigation on the yield and quality of outdoor grown tomatoes. Seed of the tomato cv. Extase were propagated in cell trays. During propagation the seedlings were fed using a stock solution containing 100 ppm nitrogen, 34 ppm phosphorous and 100 ppm potassium. The transplants were planted out on 3 December 1991 in the Karapoti Sandy Loam soil at the Plant Growth Unit, Massey University. The spacing used was 150 cm between the rows and 60 cm 2 in the row. A base maintenance dressing of Nitrophoska (12-16-10) fertilizer was applied at a rate of 500 kg per hectare banded 20 cm on either side of the row prior to planting. There were 3 conductivity treatments of 2, 4, and 6 mS cm-1 and a control treatment. A randomized complete block design was used with 4 blocks and 12 plants per plot. The 3 conductivity treatments were based on a standard greenhouse liquid feed, while the control plant received water only. Irrigation requirements were calculated based on a crop factor, area per plant and potential evapotranspiration. Conductivity treatments oommenced at the stage were 50% of the plants had commenced flowering on the first truss. Conductivity treatments were applied every 2 days for 2 hours regardless of rain, while control plants were irrigated with tap water when soil moisture deficits exceeded 28 mm day-1 except when rainfall immediately followed the scheduled irrigation. ii Plants were trained to 2 stems. The second stem was produced from the leaf axil immediately below the first inflorescence. The Otaki system of training and supporting tomato plants was used with the first support attached 25 days after planting and thereafter every 30 cm. Plants were delateraled regularly and stopped by removing the terminal buds at 2 metres high. Leaf analysis was carried out on 2 occasions, 30 and 55 days after planting, while the conductivity of the soil solution was determined at final harvest. Yield data was collected for each truss on a per stem basis per plant. Fruit were weighed individually and also size graded to the accepted commercial standard. From these data the number and weight of marketable and reject fruits were determined. Fruit samples were taken for 6 consecutive weekly harvests for compositional analysis. Firmness, total solids, titratable acidity and total soluble solids were measured from sample fruits from each treatment. Increasing the conductivity of the liquid feed increased the concentration of nitrogen and potassium in the leaves 30 days after planting, while phosphorous and magnesium were not affected by the treatments. Calcium fell with each increase in conductivity. At the reproductive stage (55 days after planting) the nitrogen, phosphorous and potassium content fell with increasing conductivity over the range of control to 4 mS cm-1 • Calcium and magnesium content also fell with increasing conductivity of the liquid feed. The conductivity of the soil solution increased as the conductivity of the liquid feed increased. As the distance from the dripper increased the conductivity of soil solution decreased. iii Tomato plants in this study supported an average of 13 trusses. There were 18 harvests where fruits were harvested at a commercial acceptable stage of maturity and a 19th harvest was used to remove all the remaining fruit on the plant. The main stem carried approximately 65% of the fruit load. Conductivity treatments had no effect on the number and weight of fruit of individual trusses on the main stem except for the 4 mS cm-1 treatment which had a higher number and yield of fruit in the third truss. No explanation can be offered for this effect. There were no differences between treatments in the number or yield of fruit per truss on the lateral .stem. Neither the number or yield of marketable fruit or the total number or total yield of fruit at final harvest were affected by the conductivity treatments. There was however a trend for yield to decrease with the 6 mS cm-1 treatment. It is possible that if the experiment had been continued for a longer period a treatment effect on the number and yield of fruit may have been obtained. It was suggested that the heavy rain experienced during the experiment may have delayed the occurrence of a yield reduction. Although there was no significant effect of conductivity on fruit size, the number of fruit in the two largest size grades tended to be highest for the control plants, while the 6 mS cm-1 treatment had the smallest number of fruit in these size grades. This is further evidence that the conductivity treatments tended to have an effect on fruit size and thus yield. The main cause for fruits to be rejected was due to fruit cracking, which usually occurred when harvesting preceded heavy rainfall. iv The occurrence of blossom end rot was low since both rainfall and the regular application of liquid feeds did not place the plants under a fluctuating moisture stress. Overall there were very few rejects. The conductivity treatments increased titratable acidity above that of control plants, but there were no difference between the conductivity treatments. Over time titratable acidity of the fruit declined and this may have been associated with either a seasonal effect or the position of the fruit on the stem. Total solids was increased as the concentration of the liquid feeds increased. The percentage of total solids allocated to structural material fell as the concentration of the liquid feed increased. This suggests that the increase in the total solids was due to an increase in the soluble solid component. There was no effect of conductivity on fruit firmness, however firmness fell from an initial value at harvest 1. Total soluble solids of the fruit increased with each increase in conductivity. Over time the trend was for soluble solids to fall slightly up to harvest 5 with a marked decline occurring at harvest 6. As improvements in fruit flavour are associated with increases in titratable acidity, total solids and total soluble solids the conductivity treatments used in this experiment were successful in improving this aspect of fruit quality. This was achieved without any decrease in yield. As suggested however, a trade off between quality and yield may have occurred if the experiment had been continued for a longer period of time. V This research suggests that the use of trickle irrigation to supply high conductivity liquid feeds to field grown tomatoes has the potential to significantly improve fruit flavour. vi ACKNOWLEDGEMENTS Writing this thesis is an enormous task. As others know well, to bring a thesis from an image to reality requires a sustaining force. For me this had been achieved through the combination of prayers and graces of good health from our Almighty and support from those who are involved at various stages of the study. My indebtedness goes to Dr. Keith James Fisher, my chief supervisor for his expertise, guidance and constructive suggestions. He was very generous and patient spending much time editing, reviewing and proofreading the manuscript. I am grateful to Dr. Michael A. Nichols my second supervisor for his confidence and camaraderie during the entire course of this work. He was very kind and helpful looking after most of my statistical analysis. I also wish to express my heartfelt thanks and gratitude to the staff of Plant Growth Unit for their unceasing support during my experimental work. It also gives me a great pleasure to record my appreciation to the New Zealand government through the Ministry of External Relations and Trade for financing my study at Massey University . My profound gratitude is also extended to the Executive Board of Romblon State College for giving me the opportunity to pursue my postgraduate degree under the Overseas Development Assistance Programme. vii My special thanks to fellow ROSCOFEANS for wishing me good luck and success in my studies. I hope I have come close to living up to their expectations. The support and encouragement I received from all Filipino students at Massey University had been of paramount importance and to all of them I wish to express my sincere thanks. The help and moral support of the Carambas and Baker families during my stay in New Zealand is here most gratefully acknowledged. My special thanks to all staff and graduate students in Plant Science (Horticulture) for their friendship, humour and help to unravel some of the mysteries in computers. My heartfelt thanks are due to my parents, brothers and sisters, in-laws and relatives for their help in many ways. Finally, my deepest gratitude to my wonderful wife Eden and son Jake for their moral support and encouragement. Their love and patient understanding are the backbone of my success. To anyone whose help during my study I have failed to acknowledge, please forgive the oversight and accept my heartfelt thanks. TABLE OF CONTENTS ABSTRACT ACKNOWLEDGEMENTS TABLE OF CONTENTS LIST OF TABLES LIST OF FIGURES LIST OF PLATES LIST OF APPENDICES GENERAL INTRODUCTION CHAPTER 1. Review of Literature 1.1 Introduction 1.2 Plant Nutrition 1.2.1 Response of Tomato Plants to Essential Nutrients 1.2.1.1 Nitrogen 1.2.1.2 Phosphorous 1.2.1.3 Potassium 1.2.1.4 Calcium 1.2.1.5 Magnesium 1.2.2 The Effect of Salinity on the Vegetative Growth of TomeJo 1.3 Flower Development 1.3.1 Introduction viii Page vi viii xii xiii xiv xv 1 3 3 4 4 4 6 6 7 9 10 11 11 ix 1.3.2 Flower Formation 12 1 .3.3 The Influence of Environment on Flower Formation 13 1 .3.4 Pollen Production and Development 16 1.4 Fruit Development 19 1.4.1 Introduction 19 1.4.2 Physical Changes in Developing Tomato Fruit 19 1.4.3 Factors Affecting Fruit Growth 23 1.4.4 Maturity and Ripening of Tomatoes 30 1.4.4.1 Structure of Tomato Fruit 30 1.4.4.2 Maturation and Ripening 32 1.5 Quality of Tomatoes 36 1.5.1. Introduction 36 1.5.2 The Composition of Ripening Tomatoes 37 1.5.3 Flavour as a Quality Attribute of Tomato 37 1.5.4 Sugar as a Component of Flavour 39 1.5.5 Organic Acids as a Component of Flavour 41 1.5.6 Aromatic Volatiles 43 1.5.7 Appearance as a Quality Attribute of Tomato 44 1.5.8 Firmness and Shelf Life of Tomato Fruit 45 CHAPTER 2. The Effects of High Conductivity Liquid Feeds on the Yield and Quality of Outdoor Grown Tomatoes 47 2.1 Introduction 47 2.2 Materials and Methods 48 2.2.1 Plant Propagation and Transplanting 48 X 2.2.2 Land Preparation 49 2.2.3 Base Fertilizer Application and Transplanting 49 2.2.4 Treatments and Experimental Design 50 2.2.5 Trickle Irrigation System 51 2.2.6 Preparation of Liquid Feed 52 2.2.7 Irrigation Schedule 54 2.2.8 Conductivity Treatments 55 2.2.9 Plant Training and Protection 55 2.2.10 Harvesting and Grading 56 2.2.11 Assessment of Fruit Quality 57 2.2.11.1 Fruit Firmness 58 2.2.11.2 Reducing Sugars 58 2.2.11.3 Titratable Acids 58 2.2.11.4 Total Solids 59 2.2.12 Leaf Analysis 59 2.2.13 Soil Conductivity and Percent Soluble Salts 59 2.2.14 Statistical Analysis 60 2.3 Results 61 2.3.1 Nutrient Content of the Leaves 61 2.3.2 Soil conductivity and Percent of Soluble Salts 62 2.3.3 Fruit Quality Characteristics 64 2.3.4 Total Number and Yield of Fruit 71 2.3.5 Marketable Fruit 72 2.3.5.1 Number and Yield of Marketable Fruits 72 2.3.5.2 Distribution of Number and Yield of Marketable Fruit 73 2.3.5.3 Size of Marketable Fruit 76 2.3.5.4 Reject Fruit 2.4 Discussion 2.4.1 Nutrient Content of the Leaves 2.4.1.1 Nitrogen 2.4.1.2 Phosphorous 2.4.1.3 Potassium 2.4.1.4 Calcium 2.4.1.5 Magnesium 2.4.2 Conductivity of Soil Solution 2.4.3 Fruit Quality Attributes 2.4.3.1 Acidity 2.4.3.2 Total Solids 2.4.3.3 Fruit Firmness 2.4.3.4 Total Soluble Solids 2.4.4 Total Number and Yield of Fruit 2.4.5 Marketable Fruit 2.4.5.1 Size of Marketable Fruits 2.4.5.2 Reject Fruit CHAPTER 3 SUMMARY REFERENCES PLATES APPENDICES xi 77 78 78 78 80 80 82 82 83 84 84 85 87 88 89 90 90 91 92 94 107 111 xii LIST OF TABLES Page Table 1. MAF Soil Test and Target Levels 50 Table 2. Standard Greenhouse Tomato Stock Solution and Resulting Liquid Feed 52 Table 3. Determination of Stock Solution (1:100) dilution Required for 3 Conductivity Treatments 54 Table 4. Nutrient Analysis of the Youngest Mature Leaf 30 Days after Planting 61 Table 5. Nutrient Analysis of the Youngest Mature Leaf Days after Planting 62 Table 6. Percent Soluble Salts 63 Table 7. Conductivity of Soil Solution ( mS cm-1l 64 Table 8. Fruit Quality Attributes in Response to Treatments 65 Table 9. Quality Attributes of Tomato During Harvest 65 Table 10. Total Fruit Number and Weight of Fruit per Plant (Harvest 1-18) 71 Table 11. Fruit Number per Plant (Harvest 1-19) 72 Table 12. Marketable Number and Weight of Fruit per Plant 73 Table 13. Numbe; of Marketable Fruit on the Main Stem per Plant 74 Table 14. Weight of Marketable Fruit on the Lateral Stem (kg per plant) 74 Table 15. Weight of Marketable Fruit on the Main Stem (kg per plant) 75 Table 16. Weight of Marketable Fruit on the Lateral Stem (kg per plant) 75 Table 17. Size of Marketable Fruits (New Zealand Fresh Tomato Industry) 76 Table 18. Number and Weight (kg) of Reject Fruit per Plant 77 xiii LIST OF FIGURES Page Figure 1. Relationship Between Dilution Rate and Conductivity 53 Figure 2. Effect of Salinity on Titratable Acids During Harvest 67 Figure 3. Effect of Salinity on Accumulated Total Solids by Weekly Harvest 68 Figure 4. Effect of Salinity on Fruit Firmness by Weekly Harvest 69 Figure 5. Effect of Salinity on the Percentage of Total Soluble Solids by Weekly Harvest 70 LIST OF PLATES Plate 1. Grading Chart Plate 2. Set-up for Weighing the Fruit Plate 3. Tomato Root System xiv Page 108 109 110 LIST OF APPENDICES Appendix 1. Growing Media Appendix 2. The Otaki System of Tomato Fencing Appendix 3. Experimental Design Appendix 4. Soil Moisture Deficit Appendix 5. Spraying Programme Appendix 6. Formula for Determining the Acidity of the Fruit xv Page 111 112 113 114 115 116 1 GENERAL INTRODUCTION The tomato is a highly valued outdoor crop in New Zealand. It is grown commercially whenever environmental conditions permit an economic yield to be obtained. Consumers continue to use large amounts of this product, but increasingly there has been publicity which suggest that tomatoes are not what they used to be. This is so because plant breeders and producers in their attempt to produce high yields have, according to a widely held view, sacrificed fruit quality. Up until recently the important quality attributes of the tomato were perceived to include size, shape, colour and firmness of the fruit. At the present time however consumers are increasingly suggesting that fruit flavour should be added to this list. It is well established that the quality of tomato fruit can vary in response to both genetic and environmental factors. It is likely that tomato breeding programs for both greenhouse and outdoor tomatoes will now start to consider improvements in fruit flavour as an important goal. Research has established that there is a close relationship between acid and sugar levels in tomato fruit and fruit flavour. Recent developments with greenhouse tomatoes has shown that increases in titratable acidity and total soluble solids can be achieved by the use of high conductivity liquid feeds. These improvements in quality have been associated with some loss of yield. 2 Drip irrigation refers to the application of irrigation water and fertilizer nutrients through small emitters placed on or in the soil near the plants. One of the advantages of drip irrigation is its ability to conserve water and fertilizer compared with other irrigation and fertilizer systems. With drip irrigation water is not applied to plant foliage maintaining drier plants and reducing susceptibility to disease outbreak with an associated reduction in the need for fungicides. Drip irrigation has been widely used with field vegetable crops in the USA. The objective of the research discussed in this thesis was to use drip irrigation to examine the potential of using high conductivity liquid feeds to improve the flavour of field grown fenced tomatoes. Of particular interest was the likely trade off between fruit quality and yield. 1.1 Introduction CHAPTER 1 REVIEW OF LITERATURE 3 Production of tomatoes has dramatically increased due to the increasing popularity of the commodity both for fresh consumption and processing. Despite the large improvement in yield that has taken place over the years, fruit quality in terms of composition has received more lip-service than the actual research effort to improve it. Consumers are becomingly concerned about the quality of the fruit they buy. As we begin to understand more of the quality attributes that consumers demand of this product, there is a necessity to develop production technologies appropriate to meet this demand without sacrificing marketable yield. It is a general observation that an important quality problem with some cultivars is the lack of flavour. This may be alleviated by means of the production system used. Part of the tomatoes success as a crop is attributed to its ability to acclimate to different environments (Rick, 1978). There are numerous environmental factors that determine wether the tomato plant live up to its potential. According to Brice (1978), tomato plants require major elements and trace elements to maintain normal growth and development. 4 Adams (1986) and Stanley and Geraldson (1991) suggest that in orderto achieve high yields and good quality fruits, proper nutritional conditions must be maintained. Regardless of the system used in the application of nutrients, the primary concern must be to provide tomatoes with the nutrients they require at each stage of growth (Stanley and Geraldson, 1991 ). 1.2 Plant Nutrition 1.2.1 Responses of Tomato Plant to Essential Nutrients 1.2.1.1 Nitrogen Nitrogen is readily absorbed by plants and easily leached from the soil. Winsor et al. (1967) reported that nitrogen fertilizer should be applied with care since excessive nitrogen is detrimental particularly during the reproductive stage. The response of tomatoes to nitrogen application, as with other plants, varies with the source (Adams, 1986), water availability, mode of application and how will the plant is supplied with other nutrients (Mengel and Kirkby, 1987). Pill and Lambeth (1977) reported that when tomato plants received NO3-N the fresh and dry weight of above ground parts increased comparably with NH4-N. The observation of Mengel and Kirkby (1987) regarding the response of various plants to NH4-N and NO3-N was that when plants received NH4-N, the uptake takes place best in a neutral medium, but was depressed as the pH fell. 5 Absorption of NO3-N on the other hand is more rapid at low pH. Uptake of nitrogen is also temperature dependent. The most important difference between NO3-N and NH4-N was that NH4-N is absorbed more readily at lower temperatures than NO3-N. It was further suggested that since photosynthesis in young tomato seedlings was low during winter it maybe safer to apply NO3-N than NH4-N. This is because plants with low carbohydrates often show toxicity when NH4-N is high. In the field fertilizer can be applied banded, broadcast or injected as in trickle irrigation. In order to stabilise nitrogen removal from soil be it as plant product, leaching or denitrification, it must be supplied in adequate amounts to the soil. During high rainfall or excessive irrigation nitrogen content around the roots is reduced particularly NO3-N, which does not react with the soil (Bar-Yosef and Sheikholslami, 1976) and NO3-N will then move with water below the root zone (Goldberg et al., 1971 ). The leaching problem with nitrogen can be alleviated by the introduction of trickle irrigation. According to Elfving (1982) following the application of the fertilizer through trickle irrigation roots of plants will have easy access to nitrogen. The emission ports are directed towards the base of the plant, and precise application of nitrogen in the plant rooting zone is possible (Stanley and Geraldson, 1991 ). By this means, nitrogen content in the soil at any one time will result from an equilibrium between addition by trickle irrigation and removal by plants plus any losses from leaching or denitrification. 6 1.2.1.2 Phosphorous Phosphorous concentration in soil solution is very dilute. It is used less by tomatoes than nitrogen and potassium, but at a steady rate throughout the life of the crop (Brice, 1978). When plants have high demand for phosphorous, the rate of absorption by the roots is high and the soil solution in the direct root vicinity is depleted of phosphorous (Olsen and Watanabe, 1970). Phosphorous in contrast to nitrogen is readily fixed in many soils (Kalkafi and Bar-Yosef, 1980). Thus, if the fertilizer is not mechanically worked deeply into the soil, most phosphorous will remain near the surface where availability will be limited. With frequent irrigation total phosphorous uptake was increased (Hedge and Srinivas, 1990). This was enhanced by movement of phosphorous towards root by diffusion, however its availability depends on soil water status. When soils are deficient in phosphorous, tomatoes become dwarfed and spindly with purple veins and leaf stalks. This deficiency normally occurs when tomato plants are grown in soils with a high capacity to fix phosphorous (Adams, 1986; Brice, 1978). 1.2.1.3 Potassium Of the macronutrients essential for growth and development of tomatoes, potassium is the most unusual since it has no direct contribution to the cellular structure of the plant (Adams, 1986). The function of potassium appears to be that of a regulator for many of the metabolic processes in cells including protein synthesis. 7 Potassium is more readily exhausted than many other nutrients. Brice (1978) estimated that tome.to plants over a season can utilize 680 kg of potassium per hectare twice as much as nitrogen. Potassium has considerable influence on fruit quality and yield (Adams and Grimmett, 1986). Winsor (1979) and Adams et al. (1978) showed that although low levels of potassium is sufficient for yield, higher levels have been shown to improve all aspects of fruit quality. Dry matter and titratable acid of the fruit generally increase with an increase in potassium level (Davies and Winsor, 1967). Winsor et al. (1961) reported that potassium improves fruit shape while Shafshak and Winsor (1964) showed that fruit firmness was also improved. Potassium deficiency is most likely to occur on heavy clay soils or on light sandy soils inadequately supplied with potassium (Adams, 1986). Symptoms exhibited by tomato plants grown in potassium deficient substrate include yellowing of the leaf margin, uneven ripening, hollowness, irregular shapes, softness and insipid flavours due to lack of acidity (Brice, 1978). 1.2.1.4 Calcium The potential uptake of calcium is very much lower compared to potassium despite its concentration in the soil which is ten times higher than that of potassium (Mengel and Kirkby, 1987). The low potential uptake of calcium according to Ferguson et al. (unpublished) arise because calcium can be absorbed only by young roots in which the endodermis are still unsuberized. 8 Apart from this uptake mechanism, the presence of other cations which are easily absorb and translocated within the plant tend to depress the uptake of calcium (Mengel and Kirkby, 1987). This partly illustrates that although soils hold comparable amounts of calcium its concentration in the plant is still low due to its controlled uptake. Ferguson et al. (unpublished) recommended that levels of other cations should be kept comparably low but above deficiency levels if the site is known to give rise to plants with calcium deficiency problems. Maynard (1991) added that apart from cation restriction, calcium uptake can also be depressed by salinity of the soil solution, low temperature, dry soil and high humidity. Calcium uptake appear mainly to be a passive process. Calcium that is absorbed by the plant roots is distributed throughout the plant principally along with water. In tomatoes once calcium has been assimilated in the leaves, it can no longer be translocated to actively growing parts with low transpiration surface (Adams, 1986). Calcium plays a critical role in maintaining cell membrane integrity. Since actively growing parts of the plant are deprived of their calcium requirement they are the first to exhibit calcium deficiency. Maynard (1991) reported that tomato plant deficient in calcium exhibit undeveloped leaves at the growing points, interveinal yellowing in young leaves and blossom end rot in the fruit (Adams, 1986; Brice, 1978). 9 1.2.1.5 Magnesium Magnesium is more abundant in soils than potassium, but its uptake rate is comparably much slower than potassium. Magnesium transport is passive moving against the electrochemical gradient (Mengel and Kirkby, 1987). Adams (1986) clarified that in tomato plants magnesium acts as an activator for certain enzymic reactions. Potassium, NH4 and calcium may play a primary role in magnesium uptake. This competition can lead to magnesium deficiency in tomato plants. Research work reported by Brice (1978) show that the average magnesium uptake over a season is 290 kg ha-1. Although high levels of potassium often depress total magnesium uptake, increase potassium supply affects magnesium content of different plant organs to a varying extent. For example, increasing potassium supply reduced magnesium content of tomato leaves and roots respectively, but the magnesium content in fruit according to Viro as cited by Mengel and Kirkby (1987), can be increased by higher levels of potassium in the nutrient solution. Magnesium is very mobile in the phloem and can be translocated from older to younger leaves. Since developing fruits are highly dependent on the phloem for their mineral supply they are thus higher in potassium and magnesium. Magnesium inadequacy in tomatoes is usually due to the plants failure to assimilate magnesium from the soil (Brice, 1978). Tomato plants deficient in magnesium show interveinal yellowing on the lower leaves and gradually which 10 spreads to all foliage. The disorder is very prevalent when plants are carrying their heaviest fruit load. 1.2.2 The Effect of Salinity on the Vegetative Growth of Tomato The development of salinity tolerance species according to Shannon et al. (1987) is an agronomic approach to the exploitation of large areas of saline soils and the efficient use of relatively abundant saline water supplies that currently have little agricultural value. As more and more land is brought into crop production through the development of irrigation, Jones (1986) has predicted the widening problem of salinity. Soils affected by salinity represent an important limitation to crop production particularly for crops that are sensitive. For tomato production, this does not pose a threat provided salinity levels do not exceed tolerable levels. Tomato plants are particularly sensitive to salinity at seedling the stage. Indications of stress in tomato were reported by Zerbi et al. (1991) as reductions in photosynthetic activity and leaf water potential which resulted later in reduced yield. Charbonneau et al. (1988) studied the effect of salinity of liquid feeds and they reported that raising the conductivity of the solution from 2 to 1 O mS cm-1 decreased linearly the shoot dry weight by 30%, a clear demonstration that raising salinity reduced vegetative growth. 11 This finding confirmed the work of Papadopoulos and Rendig (1985) who used NaCl and CaCI to raise the salinity of the nutrient base solution. Dalton and Poss (1990) suggested that salinity tolerance is controlled not only by the inherent genetic characteristics of plant, but also by some yet to be determined processes in the plant. Tomato plants that were able to withstand salinity exhibited lower growth rates and had a slightly darker green colour than the less tolerant cultivars (Rush and Epstein, 1976). 1.3 Flower Development 1.3.1 Introduction One of the major events in the life of a tomato plant is the transition from vegetative (non-flowering) state to reproductive (flowering) state. The first signs of the transition from vegetative to reproductive state, which is termed flower initiation occur at the apical stem. Once flower initiation is started, particularly in indeterminate cultivars, it continues throughout the life of the plant. Flowers whose growth and development become arrested may senesce prematurely before flower opening can occur. This pre-mature loss is known as flower abortion. 12 1.3.2 Flower Formation The vegetative period of tomato seedling lasts for about 11-31 days depending on the light conditions after pricking out (Brice, 1978). Whenever the vegetative phase is reduced, Atherton and Harris (1986) predicted early fruit production, while extended vegetative growth allows the formation of more leaves preceding the inflorescence and then delay fruiting. Hurd and Cooper (1970) suggested monitoring of leaves as an indicator of flower initiation. They suggest that 3 weeks after cotyledonary leaf expansion, most if not all tomato cultivars start to initiate flowers. This coincides with the third leaf reaching a length to just in excess of 10 mm, Inflorescence development in tomatoes require substantial amount of photoassimilate in excess of the amount required for growth. Hurd and Thornley (1974) reported that before tomato plants are set to initiate inflorescence, they have to initiate at least 6-8 leaves. This number of leaves is enough to intercept enough light needed for a sufficient production of photoassimilate for development of good quality flowers and fruits on the first truss. The hypothesis of Sachs and Hacket (1969) is in accord with the findings of Hurd and Thornley. According to this theory, the amount of photoassimilate available to the apex must reach a certain level before the transition can take place. 13 1.3.3 The Influence of Environment on Flower Formation The effects of environment can be both on flower initiation and subsequent flower development. Since flowering is determined by the appearance of visible flowers or buds, the data are sometimes insufficient to be sure which developmental stage is being affected by environmental factors. Environmental factors associated with fruit set and development in tomatoes were mostly identified under controlled environments, where such factors are subject to relatively close control. For most cultivars, the ideal day temperature for vegetative growth is about 18 to 25°C (Hussey 1973). Temperatures below this range are damaging as it reduce vegetative growth more than flower development, whereas higher temperature favour vegetative growth at the expense of flowering. Studies of Abdul and Harris (1978) show that young leaves excised from plants grown at low temperature contain low levels of diffusible gibberellins that promoted an increase in number of flowers initiated in the first truss. Leaves excised from plants exposed to high temperature contain high levels of a gibberellic acid-like diffusible substance thus giving them more advantage for attracting assimilate. Following the exposure of plants to high temperatures, Dinar et al. (1983), suggested that leaves restricted the amount of photoassimilate exported to sink organs particularly apex due to high maintenance respiration. Mofo and La Malta (1986) while studying the relationship between flowering and pre-planting 14 temperature found that once plants were exposed to low day and night temperature treatments, the number of leaves preceding the inflorescence were significantly reduced along with vegetative growth. One important result from these studies shows that inflorescence was initiated at low temperatures because the apex was able to assimilate with young leaves as their growth is restricted due to low temperatures. Koning (1991) reviewed the relationships between flowering temperature and flowering and found that at 17°C and for every 2°C rise in temperature, flowering was advanced by 0.1 truss per week. High irradiance is necessary to produce photosynthates required for flowering. As light integral increases, the number of leaves before the inflorescence decrease as does the time of truss appearance (Kinet, 1977). It appears that since the number of days to flower initiation was reduced along with a reduced number of leaves, the inflorescence is placed in a more competitive position for attracting photoassimilate over the young expanding leaves. The number of flowers in the first inflorescence tend to increase as the light integral is increased. For example, when solar radiation levels were high, flower opening occurred about 40 days after cotyledonary expansion irrespective of total radiation received (Atherton and Harris, 1986). Initial reductions in light had a greater effect than the subsequent light reductions. Flower initiation and development according to Calvert (1969) is adversely affected more than vegetative growth whenever the carbohydrate supply is insufficient. These mechanisms clearly show that when photoassimilate is limiting, the plant 15 sustainsvegetative growth in preference to other sink organs by diversion of photoassimilate from leaves and away from the inflorescence. Some cultivars are regarded as qualitative short day plants (Kinet, 1977a). Experimental evidence suggests that at the same light integral an eight hour daylength reduced the number of subtended leaves and time to initiation of the inflorescence compared to plants exposed to sixteen hour daylength. There is no questicn that carbon dioxide enrichment increases yield primarily by increasing fruit set and the final size of the fruits. Additional carbon dioxide increases the carbohydrate status of the plant through an increase in net photosynthesis. Thus from the studies conducted by Hand (1968) it was reported that carbon dioxide enrichment increases the rate of leaf initiation slightly and decreased the time to flower initiation. The effect of carbon dioxide is greater when there is an increase in light integral. Reproductive growth and fruitfulness of tomatoes relies on the correct balance between carbohydrate and fertilizer supply. Tomato plants exhibit growth retardation and flower abortion due to inorganic nutrient deficiency in the growing medium. When the macro-elements are in short supply, tomatoes tend to slow down flower induction, an indication that the plant is under nutrient stress. The size of the first inflorescence is affected by the supply of inorganic nutrients in the rooting medium when it was initiated. 16 Of the major elements supplied nitrogen is the most important. When its supply is limiting vegetative growth and flower induction is severely affected . Earliest reported effect of nitrogen on flower induction was that of Wittwer and Teubner (1957) who found that the number of flowers in the first inflorescence can be increased by increasing supply of nitrogen. The most profound effect was apparent when plants were exposed to 10-13°C when branching of inflorescence was more prevalent. Low supply of nitrogen in the growing medium increases the incidence of flower abortion. Fisher (1969) reported that low levels of nitrogen in the solution decreased flower number. The same obseNation were reported by Adams et al. (1973) when growing single truss plants. 1.3.4 Pollen Production and Development When the daytime temperature is at 20°c and under high light conditions, Picken (1984) obseNed that sexual reproduction in mother cells took place nine days before fruit set. In most instances the quality of viable pollen formed is the most important criteria in pollen production. Between 5-7 days before anthesis, damage caused by high temperature on pollen grain is even more severe than on the ovules because at this stage fruit set can be improved only by the application of untreated pollen. 17 Tomatoes are generally self pollinated (Ho and Hewitt, 1986). Mature pollen is ready for transfer at the time of anthesis while the stigma is receptive about two days previously and remains up to four days or more (Ho and Hewitt, 1986). Adherence of the pollen grains into the stigma is vital to allow germination to take place. The adherence of pollen grains into the stigma is greatly reduced when the relative humidity is 70% or if the temperature is outside the range of 17 to 24°C (van Ravestijn, 1970 cited by Ho and Hewitt, 1986). Successful transfer of pollen grains to the stigma is also dependent on the length of the style and the nearness of the stigma to the anther cone for self pollination (Rick and Dempsey, 1969). ,, Rick and Dempsey (1969) observed that tomato cultivar whose stigma is within the anther cone set fruit easily whereas in cultivars with exerted stigma or elongated style, pollination and fruit set are seriously affected. The length of the style determines the degree of insertion or exsertion of the stigma and is generally influenced by external growing conditions. Hannah and Hernandez (1984) studied the response of six cultivars grown in summer and spring conditions and found that high temperature influence exsertion while low temperature induce insertion. The observation of Munoz and Cuartero (1991) on the effects of temperature on stigma position is in agreement with the previous report. Fertilization has been first observed 18 hours after the adherence of pollen grains to the stigma and most ovules would be fertilized within 30 hours. 18 The extent of fertilization is dependent upon the number of viable pollen grains reaching the stigma, environment, physiological factors and the process of pollination and fertilization (Picken, 1984). Pollen germination is temperature dependent (Ho and Hewitt 1986). The degree of germination is greatly reduced when the temperature is below 5°C or when it exceeds 37°C. Tomato cultivars grown in the field exhibit some heat sterility specially during summer and this accounts for a reduction of yield. The most commonly accepted explanation to this has been that it was due to the abortion of pollen grains at extreme temperatures. Shelby et al. (1978) cited the study of Schaibe and reported that when night temperature reaches 32°C tomato fruit set is limited. In the same year Kuo et a/,(1979) found that most failures in tomato fruit set was due to immediate exposure of the flowers to high temperatures, but observed that if such exposure was done 5 to 8 deys after anthesis the flowers tend to tolerate the temperature level. Viability of pollen grains vary with different times of the day. When the stigma of the male sterile plants were hand pollinated with viable pollen it took 6 hours to reach the stylar end, but when pollinated early in the morning it took 12 hours (Picken, 1984). The growth rate of the pollen tube increases when the temperature is between 1 o0c and 35°C and decreases if the temperature is outside this range. 19 1.4 Fruit Development 1.4.1 Introduction Fruit set is a crucial factor that influences yield. Normal fruit growth requires two important processes namely, pollination of the flower, and the steady supply of assimilates. Undecstanding the mechanisms in these processes will enable us to manipulate and improve yield as well as the sensory qualities of the fruit (Archbold et al., 1982). Fruit number and individual fruit weight are two important components of yield. Together with sensory attributes like size and shape these are influenced by the number of seeds produced, based on the quality of viable pollen that germinates in the stigma (Picken, 1984). 1.4.2 Physical Changes in Developing Tomato Fruit Fruit growth is part of an integrated development of the plant. Fruit growth is determined by the interaction between growing conditions and morphological characters as well as physiological activities of the whole plant. It has long been recognized that the improvement of the fruit is dependent upon understanding the factors controlHng assimilate production in the leaves and the enzymatic activities that deter,·nine sink strength. The key to understanding the factors that 20 regulate fruit growth is to identify how the morphological factors respond to environment and how the metabolic processes inside the fruit interact with the rest of the plant. Fruits are reversible storage sinks as the imported assimilates are either used for growth or stored as reserves (Ho, 1988). It is indeed important to understand how the supply and the competition for assimilates for each fruit is regulated. Fruit yield in terms of dry matter production is related to assimilate supply and hence to the radiation levels intercepted by the crop. This relation however does not imply that .any increase in fruit yield is entirely determined by the improvement of photosynthesis. Fruit yield can also be improved by changing the dry matter partitioning and by manipulating the factors that determine the sink strength of the fruit (Gilford and Evans, 1981; Ho, 1988). Before fertilization, the dry matter accounts for 17% of the ovary weight. As the fruit grow, the dry matter content of the ovary as a percentage of fresh weight declines as the amount of water accumulated increases (Ho and Hewitt, 1986). The rate of dry matter accumulated declines to less than 10% after 1 0 days and 5% to 7% by day 20 and remains at this level until maturity. The rate of growth and development of tomato fruit is divided into 3 periods. First, there is a slow growth for 2 to 3 weeks when the gain in weight is less than 10% of the final weight (Ho and Hewett, 1986). The growth of the ovary ceases at anthesis and then resumes after fertilization. 21 Two days after pollination the growth of the ovary is sustained by the importation of assimilates from the nearby leaves. Thus the rate of dry matter accumulation in fruits of the same potential size will relate to the concurrent photosynthesis in the leaves. According to Ho et al. (1983), fruits accumulate as much as 30 to 50 mg of dry matter after two weeks of growth. It was subsequently observed that from twenty to twenty five days after anthesis, there was a period of rapid growth. While the growth rate increases during this period, the daily import rate of carbon diminish from 140 mg to approximately one half as the fruit increases from 20% to 90% of its carbop content. The decrease in carbon accumulated by the fruit is due to the formation of an abscission layer between the calyx and the fruit (McCollum and Skok as cited by Ho and Hewitt, 1986). When the fruit is mature green, Ho and Hewitt (1986) observed that fruits undergo slow growth marked by little addition in weight for almost 2 weeks. However intensive metabolic changes occur. The slow growth was the result of initial cell division and enlargement, while the rapid growth was entirely due to cell enlargement. Enlargement of pericarp is positively related to the auxin activity in the fruit (Asahira and Hosoki, 1977), while the shape of the fruit results from the differential growth of the ovary at the polar and equatorial dimension before anthesis. Locular cavities which vary from two and above occur as a gap in the pericarp. The locules contain seeds located in the homogenous gelatinous mass of thin walled parenchyma like cells that fill the !ocular cavities (Varga and 22 Bruisnma, 1986). These tissues appear very early and are thought to be formed by an outward growth from the placenta surrounding the seeds. The number of locules in the ovary is dependent on the flower position within the truss. Ovaries of the first flower on the first truss may have more locules than subsequent fruit. If this is the case, then the first fruit is larger than the rest, particularly those at the distal section of the truss. Since fruit size is of economic significance, increasing the locule number will enhance the size of the fruit. Introduction of genes to increase the locule number have paved the way to increasing the size of the fruit. Aside from the genes, Sawhney et al. (1971) confirmed that by applying gibberellic acid to pre-floral plants stimulated an increase in the number of carpels and locules in the ovary. In addition to these observations, the style of the treated flowers were thicker, contained more vascular bundles and there was an increase in ovary size. Reducing sugars at the onset of fruit growth account for 0.1 % of the ovary fresh weight and there was a gradual increase to 1.2% within 2 weeks and 3.5% thereafter until ripening occurred (Ho and Hewitt, 1986). Sucrose accounts for 1 % of the dry matter or in a range of 0.1 to 0.2% of the fresh weight. Although lower in percentage, it is important for fruit growth (Walker and Ho, 1979). After pollination, reducing sugars and starch increase sharply, but sucrose reduces to 1 % to 0.2% of the' fresh fruit weight within· 8 days. Although sucrose is the principal sugar imported by the fruit, its content remain low throughout. 23 According to Ho et al. (1983), the highest dry matter gain by fruits daily is 370 mg per day in the proximal fruit while dry matter increment in the distal fruit is approximately 170 mg. The major or principal part of the fruit receiving more starch per fresh weight are the locular tissues and the placental tissues. Starch which is about 1 % of the dry matter when the fruit is mature green, starts to breakdown upon reaching maximum growth (Ho et al., 1983). This concentration was further redu.ced to approximately 0.03 % of the fresh fruit weight at the time of ripening (Hobson, 1967). The breakdown of starch is associated with the accumulation of hexoses hence a correlation exists between the starch content of a mature green fruit and the reducing sugars of ripe fruits (Dinar and Stevens, 1971 ). The concentration of organic acids expressed as percentage of the fruit fresh weight was reported by Ho and Hewitt (1986) to increase during fruit development. This concentration varies according to fruit section, thus locules is more acid compared to pericarp wall and placenta. During early growth malic acid is predominant (13 %), whereas citric acid account for 25 % of the total acidity. 1.4.3 Factors Affecting Fruit Growth Tomato is a high yielding crop with fruits representing a large fraction of the total dry matter of the plant (Ho and Hewitt, 1986). 24 The reason for such a high proportion of dry matter allocated to fruit production is the high demand for sugars by the growing fruit. Developing fruit should be able to import high levels of sucrose from the leaves in order that at ripening, the fruit will contain high sugar and organic acids such that both the taste and texture of the fruit will be satisfactory. Tomato plants has a 2/5 phyllotaxy (Russell and Morris, 1983). Ho and Hewitt (1986) indicated that once the first inflorescence develops on one side of the plant, all succeeding trusses are on the same side of the stem with often 3 leaves in between trusses. The first two leaves below each truss are at angle 90° on either side of the truss, while the first leaf above the truss is on the opposite side. The assimilate supplied to the truss at flowering comes from the subtended leaves below the truss and not the leaves above the truss. At fruit initiation however, the assimilates are supplied from the leaves below the truss and the leaves above the truss (Ho and Hewitt, 1986). Tomato plants have a well structured system for distribution of assimilates and minerals to the various growing regions. The pattern of assimilate allocation to respective sinks is effective due to the arrangement of their vascular system (Russel and Morris, 1983; Yoshihiro et al., 1988). There are two main conducting tissues that sustains the growth of the fruit. The phloem, primarily conveys water and sugars from the leaves, while the xylem carries water and salts from the roots (Ho and Grimbly, 1990). In the dark, both phloem and xylem sustains the growth of the fruit, but higher growth rate in the daytime is mainly the result of an 25 increase in phloem flow bringing in sugar to the fruit. Most of the dry matter accumulated in the fruit is derived from assimilates imported from the leaves, thus fruit growth is mainly determined by the import rate of assimilates. Once carbon is fixed in a mature leaf, 20 to 30% can be exported to the developing fruits within 2 hours and approximately up to 45-50% within 2 days. The principal labile assimilate exported is in the form of sucrose making up 90% of the assimilate (Ho and Hewitt, 1986). At any one time when the supply of assimilate is limited, the fruit is sustained by both assimilates from the leaves and that remobilized from the reserves. Although the developing fruit is sustained by the leaves below the truss, transitory patterns occur depending upon the growth conditions of the fruit within the truss. Studies conducted by Bonemain according to Ho and Hewitt (1986) showed that at the peak of fruit growth large amounts of assimilate is required to sustain the needs of the developing fruits. The additional assimilate is supplied by re-mobilization of the carbon reserved in the roots and this can be re-exported within 3 hours in the form of amino or organic acids. The degree of competition for assimilate between vegetative and reproductive organs depend on their spatial and temporal relationships. In tomatoes, competition for assimilates between reproductive and vegetative organs is very localized, that is for every truss there are 3-5 leaves that act as source for assimilates (Ho and Hewitt, 1986). 26 As the developing fruits are stronger than flowers and vegetative organs, the competition for assimilates by a fruit is mainly with adjacent fruits within a truss as they derive their assimilates from the same group of leaves. The mechanisms involved in the regulation of competition between fruit is at present not fully understood. Ho (1984) elaborated that fruit growth may be determined by its inherent capacity to attract assimilate and its ability to compete with other fruits for assimilates. Most studies conducted to determine the strength of the fruit to attract assimilates involve sink size and activity as regulatory factors. Cell numbers contained in the fruit is at present considered a good measure of potential sink size. As the whole fruit mass is determined by cell number, cell size and cell content, while the final volume of the fruit is determined by the extent of cell enlargement (Coombe, 1976). Factors determining cell number in fruits have not been studied seriously. Ho (1988) stated that in tomato pericarp, the cell number appears to be associated with cambial activity of the vascular bundles . Although most cells of a fruit are derived from cell division following pollination (Mapelli et al., 1978), the difference in cell number of a fruit at various positions in the same truss already exists in the pre-anthesis of the ovary. The duration of cell division after anthesis varies from less than 2 weeks in tomatoes. Sucrose has been found directly or indirectly to be the predominant mobile assimilate in tomatoes (Walker and Ho, 1979). 27 At ripening, hexose account for about half of the fruit dry weight hence the hydrolysis of sucrose and subsequent storage of hexose may be important factors in regulating the import rate of carbon (Walker et al., 1978; Ho, 1988). If the partitioning of assimilates is motivated by the metabolic activities within the fruit, then the sink activity is a more primary determinant of sink strength than sink size. The strength of the developing fruit to attract assimilate has been thought to be the product of its dry weight. Winsor et al. (1967) studied the effect of light and carbon dioxide on fruit growth and found that al'chough the addition of carbon dioxide increased yield by increasing the number and weight of the fruit, the chemical composition of the fruit was not improved. The most accepted explanation why fruit composition was not improved, despite carbon dioxide enrichment, was that according to Ho and Grimbly (1990), although the number and size of the fruit was boosted as a result of carbon dioxide enrichment, the amount of assimilate imported by the fruit was accompanied by a similar amount of water, thus fruit weight is increased rather than the dry matter content. Both size and total soluble solid content of the fruit are influenced by the amount of light absorbed by the leaves. If light level are low during early growth the proportion of hollow fruits in the first truss is 80-100%, while dry matter content of the early yield is low. When the solar radiation is high both dry matter and sugar is high. This was demonstrated by Ho and Grimbly (1990), when they compared tomatoes imported from tropical countries with that produced in 28 Europe. Ho and Grimbly (1990) confirmed the previous observation that the amount of sugars imported by the fruit as a result of carbon dioxide enrichment is accompanied by the similar amount of water taken up by the fruit. They further explained why fruit weight increased rather than boosting the dry matter content of the fruit. The effect of light on the dry matter content of the fruit is different from the effect of carbon dioxide. Increases in light levels is accompanied by increases in temperature causing slight water stress. The fruit in return benefits from water stress by importing more sugars as the result of increased photosynthesis. High amount of light absorbed, slight water stress and a large leaf area are possible effective ways in increasing the sugar content of fruit. An efficient way of improving the quality of the fruit is the use of saline media and using the potential tolerance of tomato to salinity. As discussed by Shannon et al. (1987), the continued breeding of tomatoes to tolerate salinity has become an agronomic tool in the exploitation of saline soil not only in the field but also in controlled environment with considerable improvement in quality. Mizrahi et al. (1988) concluded that among the environmental factors responsible for the improvement of fruit quality, soil water availability and quality can readily be controlled and manipulated to increase the soluble sugar and acid concentration, thereby improving the quality of the fruit. With an increasing awareness of the effect of salinity on fruit quality, the search for materials to increase the electrical conductivity of the. solution without prejudicing the growth of the plant should have priority. In the NFT system several experiments have been conducted using different salt combinations to raise the conductivity of the solution. 29 Shalhevet and Varon (1973) added NaCl to the base nutrient solution and were able to increase the soluble solid content of the fruit by 24%. At high salinity both the uptake of water by the plant and the subsequent accumulation of water by the fruit is severely reduced, therefore the promoting effect of salinity is not on fruit density, but on the percent of fruit dry matter and sugar concentration in the fruit juice. This observation was confirmed by the results obtained by Ehret and Ho (1986), when they observed the dry matter of the fruits from salinized plants to increase ,as the conductivity of the solution increased. During rapid growth, Ehret and Ho (1986) observed that a high proportion of dry matter was stored in the fruit in the form of starch rather than sugars. The advantage of accumulating carbohydrates in fruits in the form of starch rather than hexoses according to Mitchell et al. (1991) is that it ensures continued sink activity therefore the accumulation of starch is maintained at a reduced tissue water level. These studies clearly demonstrate that when the assimilate supply is not limiting the amount of assimilate being imported by a fruit is not at all affectec;I by the water relations in,the plant. Although size of the fruit is reduced (Ho and Grimbly, 1990) the fruit have a higher percentage of dry matter. If for instance the purchase of the produce is based on the total solid content there would be no economic loss. When sugars are not limiting, fruit growth is regulated by temperature and water supply. Restricted watering and high salinity can be used to adjust fruit number, fruit size and improve taste. The effect of such reduction is not to reduce 30 phloem sap supply, but to increase the concentration of the sap without a reduction in the usual amount of sugar being imported by the fruit. Although less water is accumulated, the dry matter accumulated increase with high sugar concentration in the fruit. The effect of macro and micro elements has been discussed by Uexcull (1979). Adequate nitrogen improves fruit quality, while excessive application tend to decrease the size of the fruit, keeping quality, colour and taste. Phosphorous on the other hand in combination with potassium improves peel and pulp coloration, taste, firmness, vitamin C content and hastens maturity. Potassium tends to increase fruit size and plays a key role in the process of fruit pigmentation. It also influence acid metabolism. Inadequacy of calcium in the growing media will result to a number of physiological disorders.while magnesium deficiency can be induced by high rates of potassium or ammonium nitrogen application. 1.4.4 Maturity and Ripening of Tomato 1.4.4.1 Structure of Tomato Fruit Davies and Hobson (1981 ), botanically classified tomato fruit as a berry since the seeds are formed within the fleshy mesocarp. Tomato fruit is composed of flesh (pericarp wall, skin and pulp), placenta and !ocular tissue including seeds. Generally, less than a third of the fruit fresh ·weight is made up of pulp (Ho and Hewitt, 1986). 31 The skin is made up of 4 to 5 layers of cell under a thin cuticle. The epidermal layer is heavily cutinized on the surface. The epidermis is covered by a thin cuticle that thickens as the fruit matures ( Varga and Bruinsma, 1986 ). The cutin may surround the epidermal walls below and between the first layer of collenchyma. The importance of cutin is enhanced resistance of the fruit to cracking during ripening. This is very important particularly when tomatoes are grown purposely for processing and mechanically harvested and transported for long distances (Voisey et al., 1970). Vascular strands radiate from the stem end of the fruit to the pericarp down to the columella and to the blossom end. Cross connection in the pericarp are more common in the distal half of the fruit. A major vascular bundle, the centre line of each carpel, is developed extending from the stem scar to the blossom end of the fruit (Ho and Hewitt, 1986). In green fruits, the epidermal cells tend to have less starch than the inner parenchymatous cells. Most of the cell division in the pericarp takes place during the first week after anthesis, further division occur during the second week (Asahira et al., 1965 cited by Ho and Hewitt 1986). The number of cell layers increase from 8 to 1 O during this period however cell development was observed throughout fruit development. In mature fruits, the placental tissue is firm, but as it approaches maturity, the walls of these parenchymatous cells become thin and wavy and contain large but diminishing numbers of starch grains (Grierson and Kader, 1986). The appearance of jelly like material in the !ocular cavity provides an excellent 32 criterion whereby fruit may be designated mature green. The placental tissue at incipient ripening appears pink in colour. Two major groups of pigment in tomato fruits are chlorophyll and carotenoid. Prior to ripening chloroplasts lose starch and chlorophyll from the grana, followed by the production of osmophilic globules and the formation of lycopene within the swollen granum compartment. 1.4.4.2 Maturation and Ripening The fruit of tomato requires approximately 40-60 days from flowering to full ripeness. After the initial phase of slow growth, Rick (1978) observed that the fruit undergo major increase up to full size in about half of the ripening period. As the fruit matures, cells in the pericarp become very large, thin walled and the parenchymatous cells around the seeds are reduced to semi-liquid immediately prior to ripening (Ho and Hewitt, 1986). The starch content reaches a peak halfway through development, when it makes up about 20% of the dry matter. This figure falls to 1 % at the mature green stage and is followed by a declining growth rate during which maturation and ripening usually occur. During this period, the fruit (Varga and Bruinsma, 1986) stores large amounts of reserve materials accumulated either through synthesis of sugars or elaborated in the nearby source leaves. Early ripening is marked by the completion of starch hydrolysis at which point reducing sugars are present in maximum concentration. At almost the same time malic acid, which was initially an important organic acid decreased gradually after 33 fruit maturity and is replaced by citric acid. The turnover in acid concentration in favour of citric acid causes the ratio of malate to citrate acid to fall during the later stages of maturity. Hobson and Harman (1986) elaborated that at maturity, the sugar content in the fruit, in particular reducing sugars, tends to be more concentrated in the !ocular walls while acids are more concentrated in the locular cavities. An increase in moisture content was also observed in the fruit, which accounts for 90-95% of fresh weight as the fruit develops. Aside from compositional changes, a series of structural changes are known to occur. Following a decline in growth rate, Bathgate et al. (1985) noticed gradual structural changes including membrane disintegration, chlorophyll breakdown and starch disappearance. Gradual chlorophyll degradation at this stage results in the whitening of the fruit which serves as a useful determinant of ripening. The following week is marked by destruction of the remaining chlorophyll when Harris and Spurr (1969) observed the transformation from chloroplast to chromoplast. During this transition, there was an increase in carotenoid content and synthesis of lycopene in the cytoplasm. Grierson (1987) noted that the mechanism of change was structured so that it took place at the same time that other ripening processes such as cell wall degradation, tissue softening and accumulation of sugars and organic acids were occurring. Ripening of tomato fruit involves a number of chemical and physical changes which convert the fruit from a relatively inedible state to one of optimal quality (Babitt et al., 1973). It is also a complex developmental process involving 34 changes in the biochemistry, physiology and gene expression of the fruit and an active process characterized by changes in all cellular compartments as well (Smith et al., 1989). It has been suggested by Saccher (1973) that the loss of cell membrane integrity, removal of permeability barrier and increase in free space serve as major factors in ripening and senescence. One of the early events that heralds the onset of ripening is the beginning of the respiratory climacteric, the rise in breakdown of proliferated cells that surround the seeds in the locules (Rabinowitch et al. 1975), the partial solubilization of the cell wall components leading to softening of the texture (Speirs and Brady, 1991 ), alterations in metabolism and composition of all major cell compartments (Grierson, 1987) and progressive changes in colour, texture, flavour and aroma in each tissue (Brecht, 1987). All these changes are highly synchronized appearing in close succession in a variety of distinct processes during the relatively short period which fruit ripens. The mechanisms by which fruit ripening changes are initiated and synchronized remains largely unknown. It therefore seems appropriate to look into factors that triggers the initiation and coordination of these changes. Essential to the initiation of ripening is the resistance of the fruit tissues to the effects of hormones that would trigger metabolic changes within the cells. Resistance of tissues to ripening hormones is a feature of most unripe tissues (Hobson and Harman, 1986). Once their resistance has been overcome by ripening hormones their evolution gradually rise from a low basal level. 35 The rate at which tissue sensitivity towards the ripening hormone occur determines the onset of ripening. One important and much studied process occurring during fruit ripening is softening. This involves changes in the structure and solubility of the fruit cell wall. The processes involved in fruit ripening has been extensively studied. The main issue under investigation here is the occurrence of ripening enzymes that eventually lead to metabolic degradation of fruit cells. Previous efforts to determine the enzyme that initiated ripening was made by Sacher (1973). Accordingly, there are numerous enzymes that have been shown to change their activities as the fruit ripens, and some of these show clear physiological relevance. For example, when Brecht (1985) measured the concentration of various enzymes in tomato fruit tissues, he found that during maturation the concentration of ACC increased in the locule tissue of green fruit coincident with gel formation. Brecht (1987) clarified that !ocular tissue was derived from the placenta of the fruit then overgrowing the seeds, which later on fill the locules, but not fusing with either the seeds or pericarp. At maturity, the cell walls become progressively thin and fragile eventually rupturing to produce the typical liquid jelly like consistency of mature ripe fruit. The activity of ACC synthase is not only ' centred on the !ocular tissues but is spread out to nearby tissues like columella and placenta, whe(e synthase begins to increase at the mature green stage (Brecht, 1985). Similar increases also occur on radial and outer pericarp following the breaker stage. 36 ACC and ACC synthase continue to increase in columella and pericarp tissues beyond the breaker stage while levelling off (ACC) or falling (ACC synthase) in the gel as the tissue is degraded. The expansion and separation of cells and the cell wall fragments released at this stage, may act as elicitors of ethylene production (Brecht, 1985). The locular contents of tomatoes contributed 20% to the total fresh weight of the fruit thus a small increase in ethylene production observed in the whole fruit at the mature green stage probably represent a significant increase in the locule tissues rate of ethylene production. The relationship between ethylene production rate and maturity stage is a good index to determine physiological maturity of the fruit. 1.5 Quality of Tomatoes 1.5.1 Introduction Tomato quality has received attention only after the improvement of yield has been achieved. Consumer preference for fresh tomatoes is no longer limited to how they appear, but they are also demanding quality attributes based on flavour and nutritional value. It is generally accepted that ripe tomatoes purchased in supermarkets nowadays lack the desired aroma and flavour that there used to be when fruits were vine ripened. U.S.D.A. legislation regarding product information requires the inclusion of nutritional value of the product. This move will change the purchasing attitude 37 of consumers as they can compare product composition. The impact will not end where the product is being sold, instead growers will be obliged to do their part by changing their production practices so that their product will be competitive on the market. 1.5.2 The Composition of Ripening Tomatoes Tomato fruits ripening either on or off the vine show compositional changes. It is at maturity that the ultimate quality of the fruit can be judged based on the perception of consumers. Tomato fruits during ripening contain a wide range of compounds and show considerable variation in composition and structure. The presence of many compounds that constitute the flavour of the fruit is the result of interactions with season, nutrition, environmental and physiological status of the fruit. The composition of present day tomato varieties is the result of combined efforts of research work to determine the factors that comprise fruit flavour in particular sugar, acid, and volatile compounds. 1.5.3 Flavour as a Quality Attribute of Tomato Flavour is one of the most important constituents of quality, but at present has little effect on the initial sale of the product due to our failure to identify its 38 components. Tomato fruit are sensed differently by different individual and often impressions are affected by other non-flavour quality attributes. The unique flavour of tomato is a composite of taste smell and touch or mouth feel (Bezar, 1989) and is a product of sugars, acids and aromatic volatiles contributing to 60 % of the dry matter Hobson (1988). If the proportion of sugar in the fruit is low, the consumer will react to their sharp and biting taste, but if the fruit acid is low the unbalanced sweetness can be disturbing to the taste. Weight for weight acids affects the taste more than the sugars. Of the sugars that are used to balance acidity fructose has more sweetening power than glucose due to its higher concentration. Volatile compounds are not only important to aroma, but also to the overall flavour of tomatoes. Frenkel and Jen (1989), reported research work and concluded that although several hundred aromatic volatiles have been catalogued, there are still new compounds being discovered and added to the existing ones as research for other volatile compounds continue. Only a limited number of these are important to the aroma of the tomato fruit. Improving the sugar and acid ratio of the fruit requires effective combination of factors that influence the chemical composition of the fruit. Studies conducted by Ehret and Ho (1986) and Mizrahi et al. (1988) revealed that among the environmental factors that influence the quality of the fruit, nutrient and water availability and quality can be readily controlled and manipulated to increase the total soluble solids and acid c9ncentration. Mizrahi (1982) reported that increasing the concentration of the liquid feed increases the total soluble solids and acid of the 39 fruit. By imposing a range of salinity regimes, water uptake is regulated thus increasing the concentration of mineral ions in the fruit. The concentration of cations in the fruit varies with the kind of salts used to raise the conductivity of the nutrient base solution. Thus, when Ehret' and Ho (1986) imposed a range of salinity using potassium nitrate or calcium nitrate, potassium becomes the predominant cation in the fruit. An important factor in understanding the determinants of flavour is to know how to balance the acid and sugar content of the fruit. Bezar (1989), pointed out that fruit walls contain at least 20 % more sugar than the juice around the seeds, while the juice contain more acid than the pulpy part of the fruit. Hobson (1988) elaborated that if consumers wanted to derive a biting taste they should choose those fruits with bigger locular contents in contrast to pulpy fruits which gives a sweeter taste. 1.5.4 Sugar as a Component of Flavour After water, sugars form the next most important constituent of the tomato fruit contributing almost 50% of the total dry matter of the fruit of commercially grown varieties (Winsor, 1966). Ripe fruit contain free sugars in the form of reducing sugars consisting of glucose and fructose in approximately equal amounts, but usually with a preponderance of fructose at late stage of ripeness (de Bruyn et a/., 1971; Davies and Hobson, 1981 ). On equal weight basis fructose is sweeter than glucose. 40 Hobson and Kilby (1985) and Baldwin et al., (1991) indicated that tomato fruit of normal size contain very little sucrose. Initially, Davies and Kempton (1975) reported that as the fruit develops there is a progressive increase in both glucose and fructose concentration and then they show a dramatic increase during the stage of cell enlargement where the ratio of glucose to fructose decreases rapidly. The pronounce rise in sugar concentration in the fruit coincides with the first appearance of yellow pigment in the walls accompanied by changes occurring in the whole fruit, !ocular juices and pericarp. Environmental and production practices greatly influence the sugar concentration of tomato fruits. Davies and Hobson (1981) pointed out that of the known environmental factors affecting the composition of the fruit it is probable light level has the most profound effect on sugar concentration. Thus seasonal trends in sugar levels reflect differences in the intensity and duration of light. Growers have limited opportunity to manipulate the sugar content of the fruit except for morphological modifications to improve light interception and agronomic practices to maintain the vigour of the plants and to increase the rate of photosynthetic production. Since fruits of most typical tomato cultivars contain 95% water the dry matter therefore constitute a very small fraction of the fruit. Ho and Grimbly (1990) estimated that the dry matter of the fruit comprises 8% minerals and the rest consists of carbon products. To increase the proportion of dry matter growers have to restrict the water uptake of the fruit by increasing the osmotic potential of the nutrient base solution and in order to maintain the osmotic 41 gradient the plants accumulating more solutes in the fruit (Mizrahi, 1982; Hobson, 1988; Adams, 1991 ). 1.5.5 Organic Acids as a Component of Flavour Organic acids are one of the major taste components of tomato fruit. The acid content of the fruit is an important characteristics of processing tomatoes. To avoid problems with thermophilic organisms, Stevens (1972) suggested that the acid content of the fruit must be high enough to provide a pH of not less than 4.4. High ph values however necessitate longer processing times increasing the difficulty of obtaining high quality product. There is a continual change in the acidity of the tomato fruit during development and maturation. The concentration of acids increase during development and reach maximum near incipient colour and then decrease until well beyond maturity (Stevens, 1972). When fruits are still immature malic acid is predominant, whilst citric acid forms only about 25% of the total acidity (Stevens, 1972). Maximum acidity during ripening coincides with the first appearance of pink colour, when malic acid concentration decreases rapidly in !ocular juices and in the outer fruit walls (Davies and Hobson, 1981 ). As a result of different rates of change during ripening, malic to citric acid ratio rapidly declines. Thus citric acid makes a greater contribution to sourness than malic because it occurs at a higher concentration (Stevens et al., 1977). Titratable acidity of the outer fruit wall is relatively low compared to that of locular contents. 42 Should there be any particular change in titratable acidity, it has always been attributed to citric acid alone or to changes in both citric and malic acids. The relationship between potassium and acidity in tomato fruits is very close and highly significant (Varga and Bruinsma, 1986). Potassium accounts for 85% of , the total cations in the fruit (Davies and Hobson 1981 ). This relationship varies according to the level of the potassium supplied and the growing conditions (Besford and Maw, 1975). At high potassium supply, the acidity increases (Davies and Winsor, 1967) and the colour and shape improves (Winsor et al., 1961 ). At low potassium supply, the growth period of a tomato fruit is shortened (Besford and Maw, 1975) and the maximum climacteric respiration is enhanced. Mengel and Viro (1974) reported that the potassium concentration affected the metabolism of the fruit. The form of nitrogen applied influences the acidity of the fruit. With ammonium nitrogen Varga and Bruinsma (1986) observed the fruit to be less acid in composition than with nitrate nitrogen. When high potassium and nitrogen is applied the combined effect is particularly favourable to fruit acidity, while magnesium and calcium have little effect. Calcium when combined with increased potassium lowers the acid content of the fruit. Tomato fruit is a valuable source of vitamin C. Typical values of Vitamin C vary from 16-25 mg/100 g fresh weight. Extensive literature reported by Hobson and Davies (1981) is no doubt a reflection of its importance to human nutrition and 43 the contribution that tomato fruit can make. Ascorbic acid content of tomato fruit varies. The wide variation is probably attributable to differences in light conditions during growth. It is not surprising to note that fruits harvested from plants grown outdoor were found to contain more ascorbic acid than fruits grown under glass. Similar effects of light on ascorbic acid content can be found on fruit positioned in the plant depending on their relative exposure to sunlight. Thus, a rise in ascorbic acid content with ascending truss position was probably due to increasing light intensity. 1.5.6 Aromatic Volatiles The demise of the tomato like flavour is not only associated with the imbalances between sugars and acid, but also due to the lack of a desirable aroma in ripe fruits. Volatile compounds responsible for the odour of tomatoes are present in a relatively small amounts. The odour of freshly picked tomatoes comes from the calyx, sepals and in the stalk rather from the fruit per se. The skin of tomato fruits has no pores or lenticels so that volatile compounds cannot escape. It is only by means of cutting the fruit that we can smell the aromatic compounds. There have been extensive studies conducted to identify the volatile compounds of fresh tomatoes, but inspite of these trials the actual quantitative concentrations has not been reported. Hobson (1988) reported that there are 400 substances that make up the odour of tomatoes, but not even one of them has a smell 44 reminiscent of a rip~ tomato. The flavour of tomatoes is a combination of aroma and the taste components modified by the texture of the tissues and its juiciness. Buttery et al., (1988) showed that the pulp and skin had the highest concentration of the volatile aroma compounds and no significant contribution related to the seeds. 1.5. 7 Appearan~e as a Quality Attribute of Tomato The quality of tomato fruits is determined among other factors by their size and shape (Favarro and Pillatti, 1987). Size and shape are quality attributes important in marketing tomatoes because it is the first attribute perceived by consumers. Size and shape is standard criterion for grading and packaging. Bangerth and Ho (1984), reported that fruits within the same truss generally differ in their final size, thus, larger fruits develop at the proximal end compared with the distal fruits. The differential growth based on their position in the truss is not entirely due to competition, because if the number of fruits is reduced, the remaining distal fruits still remain smaller. Bangerth and Ho (1984), demonstrated that the final size of the fruit can be manipulated by the induction sequence in the truss, that is by inducing the growth of the fruit in the distal end of the truss ahead of the proximal fruits, a much larger fruit can be produced. As discussed previously, ovule number influence fruit size therefore a large many seeded fruit is likely to originate from an ovary containing many ovules. According to Mizrahi et al. (1988), saline treated plants produce small sized fruits 45 as they contain less water and more soluble solids. Colour which depends primarily on the content of total carotenoids and ratio of lycopene to carotene is another quality attribute of tomato fruit. Most consumers prefer deep uniform red coloured tomatoes as a basis for selection. At the ripening stage, temperature has a great influence on the rate and extent of colour change. Lower than normal temperatures lead to an increase in beta-carotene and decrease in lycopene. When tomato fruits are exposed to temperatures between 30-40°C they remain yellow rather than red (Davies and Hobson , 1981 ). Different morphological regions of tomato fruit vary in chemical composition. Total carotenoids are found to be highest in the outer pericarp whereas carotene is highest in !ocular region. Mizrahi et al., (1988) observed that fruits from salinized plants were redder in colour than the normal fruits. The effect was intensified as the conductivity of the nutrient solution was increased. Increasing the concentration of potassium in the solution has been found by Dangler and Locasio (1990), to reduce the severity of blotchy ripening. 1.5.8 Firmness and Shelf Life of the Tomato Fruit Firmness is a quality factor that determines the shelf life of the fruit. Firm fruited cultivars show a great advantage hence fruits can be allowed to ripen on the vine until almost 100% of the fruits is ripe. This practice according to Stevens (1980) can increase the viscosity of the fruit which provides tomato products with increased consistency. Changes in fruit firmness are due to the activity of 46 softening enzymes. Since firm fruits have reduced amount of !ocular tissues, they exhibited lower rates of respiration and ethylene production during ripening. One disadvantage of firm fruited cultivars is that they contain less acid thereby lowering their acid· to sugar ratio Stevens (1977). Firmness of the fruit show variations along the morphological regions of the fruit. Adegoroye et al., (1989) showed that there is a progressive decrease in deformation from stylar end of the fruit to the calyx, whilst the shoulder part of the fruit is found to be the weakest point. Both epicarp strength, !ocular resistance and firmness decrease, while deformation and compliance increase with ripeness. Mizrahi (1988) reported a contrasting effect of salinity on the shelf life of tomato. It was observed that saline treated plants produce fruits that are slightly firmer, but deteriorated somewhat faster than the normal fruits in contrast with their other observations showing that the shelf life of the fruit was extended due to a reduce amount of water in the fruit. CHAPTER 2 THE EFFECTS OF HIGH CONDUCTIVITY LIQUID FEEDS ON THE YIELD AND QUALITY OF OUTDOOR GROWN TOMATOES 2.1 Introduction 47 High yielding good quality outdoor tomato crops are traditionally grown on fences in the Otaki district of New Zealand. Crops are harvested from summer through to late autumn and are irrigated by using overhead sprinklers. Nutrients are applied as a base dressing along with a number of side dressings. Trickle or drip irrigation has been used for some time with vegetable crops in Israel and is under investigation in the USA with an extensive range of crops (Hochmuth, 1992). It has not been used to any extent with outdoor vegetable crops in New Zealand. Research with greenhouse tomatoes has shown that fruit flavour can be improved by the use of high conductivity feeds. The effect of these treatments has been to increase the concentration of sugars and acids in the fruit (Anon, 1992). In Israel, Pasternak et al. (1986) have shown that total soluble solids of trickle irrigated tomatoes was increased by the use of saline water, while Cornish and Nguyen (1989) in Australia have shown that high soil electrical conductivity was found to increase total soluble solids comparable to that of hydroponic systems reported by Hobson (1988). 48 The following study was designed to use trickle irrigation to apply high conductivity feeds to fenced tomatoes to determine the effects of such treatments on fruit yield and quality. 2.2 Materials and Methods 2.2.1 Plant Propagation and Transplanting On 21 October 1991, seeds of the tomato cultivar Extase were sown in cell trays at one seed per cell and placed on a heated bench (24°C). The trays held 54 plants and 85 mis of growing media per cell. A 50 : 50 peat sand media was used (Appendix 1 ). At emergence the seedlings were removed from the heated bench and grown on in a greenhouse with a minimum temperature of 16°C and ventilation at 22°c. Thirty days after germination, and thereafter twice a week until the plants were ready for planting, the seedlings were fed using a stock solution containing 100 ppm N, 34 ppm P and 100 ppm K. On , 27 November, one week before transplanting, the seedlings were moved out of the glasshouse and hardened off outside. 49 2.2.2 Land Preparation The experiment was conducted during summer of 1991 on Karapoti Sandy Loam soil at the Plant Growth Unit at Massey University. The experimental block was sheltered and the land was ploughed, rotovated and levelled to prepare a planting out tilth to a depth 20 cm. One hundred and fifty soil samples from the top 15 cm were collected at random from the 225 m2 experimental area using a soil core borer and a sub-sample was submitted to the MAF Laboratory for analysis. Ten fences, 1.5 m apart and 15 m long were then erected based on the system used in the Otaki district. (Appendix 2). The post were 2 m above the ground and were placed 3. 75 m apart in a row. 2.2.3 Base Fertilizer Application and Transplanting The result of the MAF soil test and target values based on the recommendation of Clarke et al. (1986) are presented in Table 1. As the soil test data were satisfactory, a base maintenance dressing of Nitrophoska fertilizer (12-10-1 0) was applied at a rate of 500 kg per hectare before transplanting. The fertilizer was applied in band 20 cm. on either side of the row and push hoed into the soil. 50 Table 1. MAF soil test results and target levels. pH Ca p K Mg Na Soil Test 6.1 12 41 11 27 7 Target Values 5.3-6.7 10 33-45 13 15 - On 3 December 1991, forty days after germination,the plants were transplanted and watered in while the guard plants were planted a day later. The plants were in rows 1.5 cm. apart and 60 cm. apart in the row. 2.2.4 Treatments and Experimental Design There were four treatments, control and three conductivity treatments. The control treatment receive no further nutrients and was watered as required. The three conductivity treatments were 2, 4 and 6 mS cm-1 • These treatments were applied from 31 December and were based on a standard nitrogen-potassium greenhouse liquid feed (Table 2) . A randomized complete block design was used. A block consisted of 2 rows of plants with 2 plots per row (Appendix 3). The experimental plot was 7.2 m long 51 and contained ten plants with 2 guard plants at the end of each row and between adjacent plots. There were two guard rows at each side of the planting. 2.2.5 Trickle Irrigation System The trickle irrigation system used in this experiment was connected to the Palmerston North City Corporation Water Supply. A valve and pressure gauge on the main line controlled the delivery to the system. The main line was split into 4 submains to deliver the four treatments (Appendix 3). lnline diluters (Dosatron 1501) were connected to the 3 submains to supply the conductivity treatments and the fourth sub-main supplied water to the control treatment. The four sub-mains were laid across the centre of the experimental block. In each replicate lateral pipes were connected to the sub-mains according to treatment. Typhoon 25 dripper-line with a water inlet 60 cm apart were used for the conductivity treatments, while pvc pipe with microsprinklers was used for the control plants. The microsprinklers were spaced 1.2 m apart. Each microsprinkler was set to water 2 plants, one on each side. The four lateral pipes were fixed in place by tying them to wires attached to the posts. The layout of the diluters, nutrient tanks, submains and lateral pipes is detailed in Appendix 3. 52 2.2.6 Preparation of Liquid Feed The three conductivity treatments used provided conductivities of 2, 4, and 6 ms cm-1 • The liquid f~eds for these treatments were based on a standard greenhouse stock solution (Table 2). A series of test dilutions were prepared using the above stock solution to determine the right amount of the fertilizer required to be dissolved to provide conductivities of 2, 4 and 6 mS cm-1 • Figure 1 presents the relationship between the dilution rate and conductivity derived from the test solution prepared. Table 2. Standard greenhouse tomato stock solution and resulting liquid feed (Clarke etal.,1986). Composition Dilution 1 :200 Fertilizer (grams litre-1 ) N K Mg s (ppm) (ppm) (ppm) (ppm) Potassium 150 105 285 0 0 Nitrate Urea 30 57.5 0 0 0 Magnessium 50 0 0 60.75 80 Sulphate The dilutions of the standard stock solution (Table 2) that provided the desired conductivities were read off the graph (Figure 1) and then used to determine the stock solutions that when diluted 1 :, 00 would produce the three treatment conductivities (Table 3). Figure 1. Relationship between dilution rate and conductivity of liquid feed. 6-,----------------------'---- 11 :1051 5.5 - 5 E ~ ~ 4.5 ........... ~ :s: 4 u ::J "O 3.5 C 0 () 3 2.5 25 1 :46 50 Dilution rate 100 53 Table 3. Determination of stock solution (1:100) dilution required for three conductivity treatments. Dilution Potassium Urea Magnesium Treatment required (read Nitrate Sulphate (mS cm-1 ) off from graph) Amount required grams litre-1 (Dilution 1:100) 2 1 :105 142.85 28.60 47.20 4 1 :46 326.10 65.22 108.70 6 1 :29 517.50 103.50 172.50 54 Stock solution were then prepared which when diluted 1 :100 provided the treatment solutions detailed in Table 3. Since the weight of potassium nitrate required to provide the 6 mS cm-1 treatment exceeds its solubility two stock solutions each capable of supplying 3 mS cm-1 were used . 2.2.7 Irrigation Schedule Irrigation requirement of the plants was calculated based on a crop factor, area per plant and potential evapo-transpiration. Assuming that the area per plant is 50% of the true area, each plant occupied 0.45 m2 • The crop factor for greenhouse tomatoes of this size is 1.2 ( White, 1989). Thus for every millimetre of evapotranspiration, 0.54 litres of water is required by each plant. As the average potential evapotranspiration is 4mm day-1 , each plant will require 2.16 litres of water daily. The water requirements for control plants was based on soil moisture deficit. If the estimated soil moisture deficit did not exceed 28 mm per week no irrigation was applied to the control plants. 55 Information regarding soil moisture deficit were taken from rainfall data from DSIR, Grasslands Division and potential evapo-transpiration (4 mm day-1 ). Based on this information, there were only four occasions from January to April when the soil moisture deficit exceeded 28 mm. However control plants were irrigated only twice as on two of the occasions that the deficits exceeded 28 mm there was heavy rain before the irrigation could be applied (Appendix 4). Water was applied for two hours on the two occasions irrigation was supplied. 2.2.8 Conductivity Treatments The conductivity treatments were applied from 31 December 1991. At this stage setting in the first truss had commenced on 50 % of the plants. The conductivity treatments were irrigated every 2 days thereafter for a duration of two hours. This applied 3.6 L of liquid feed per day. The conductivity of the treatment solutions was monitored after irrigation at different locations using a CF meter. 2.2.9 Plant Training and Protection The Otaki system sit training and supporting tomato plants was used. This was done by erecting 4 posts ( 15 cm in diameter and 2.5 m long) in a row set 3. 75 m apart to a depth of 60 cm. End row posts were angled to take the strain once the wire supports were stretched the length of the rows. The rows were 15 meters long. 56 A pair of wire was attached to the posts at 30 cm intervals from the ground. Plants were trained between pairs of wires which were laid out as the plants required support. Plants were trained to two stems. The second or lateral stem was produced from ·the axil of the leaf immediately below the first inflorescence. The first pair of wires was attached 25 days after planting. The two stems were delateraled at regular interval and the plants were stopped by removing the top l of the plants with a: knife at two meters. A push hoe was used once every two weeks to control weeds. A regular weekly spray programme was carried out for disease and pest control. Details are presented in Appendix 5. This programme was repeated every three weeks. 2.2.10 Harvesting and Grading To provide details and information about the position on the plant were the fruits were harvested, each truss was labelled using different colour tags according to their position on the individual stems. Fruits were harvested once they reached a red orange colour. There were two harvests per week for the first two weeks and three harvests per week thereafter. Harvesting commenced on March 9 with the final harvest on May 1, 1992. Fruits were harvested on an individual truss basis for each treatment. Each fruit was weighed individually and graded according to size (diameter) with reject fruits being allocated to 5 classes . 57 Fruit sizes were classified using the grading chart recommended and issued by New Zealand Fresh Tomato Industry (Plate 1 ). Fruit were therefore graded to the following diameters; 70+, 60-70, 50-60, 40-50, 40 mm and below. These sizes were coded as grades 1-5. For fruits to be considered marketable they were free from physical damage and uniform in shape and colour. Rejected fruits were indexed as cracked, rough, blossom end rot, blotchy and diseased. Fruit weights were determined by weighing the fruit individually using a Mettler scale, interfacing with a laptop computer and printer to record the weight along with the information on fruit size and grade (Plate 2). Total yield was computed as combined weights of marketable and rejected fruit. This however did not include the fruits that abcissed from the trusses before they were ripe and those that were damage by birds, as it was difficult to identify where on the plant such fruit originated. 2.2.11 Assessment of Fruit Quality Ten sample fruits were chosen at random from the fruit harvested from each treatment plot once a week for six consecutive weekly harvests starting from March 19, 1992. · Sample fruits were of the same size and maturity with diameters ranging from 50-60 mm. Since the fruits were harvested when orange red, they were ripened at ambient room temperatures until the red ripe stage was reached before quality assessments were made. All quality assessments were carried out on the same day. 58 2.2.11.1 Fruit Firmness Firmness of sample fruits were measured by using a penetrometer. Measurements were taken from th