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SOIL WATER DEPLETION, GROWTH, PHYSIOLOGY, AND YIELD OF CARAMBOLA TREES IN KROME SOIL

SOIL WATER DEPLETION, GROWTH, PHYSIOLOGY, AND YIELD OF CARAMBOLA TREES IN KROME SOIL

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Carambola (Averrhoa carambola) is a commercially important tropical fruit tree in South Florida. Little is known about carambola water requirements or its response to limited soil water supply under subtropical conditions such as those of southern Florida. The objective of this study was to determine the effects of four levels of soil water depletion (SWD) that ranged from field capacity (FC) to the first visual signs of water stress (leaf yellowing and abscission), on leaf gas exchange, stem water potential, phenology, growth, and yield of 2-year-old, container-grown carambola trees, and 8-year-old 'Arkin' carambola trees in an orchard.
Soil water depletion treatments were determined from continuous measurement of soil water content using multisensor capacitance probes (EnviroSCAN, Sentek PTY Ltd., Kent Town, Australia). Irrigation of orchard trees was initiated when SWD reached one of the following levels (where 0% SWD = FC): 0-8% SWD, 9-11% SWD, 12-14% SWD, or 15-17% SWD. Trees in 95 L containers filled with Krome very gravelly loam soil were irrigated when SWD reached one of the following levels: 0-21% SWD, 22-31% SWD, 32-50% SWD, or 51-60% SWD.
An increase in SWD resulted in a reduction in stem water potential (ΨS), net CO2 assimilation (A), transpiration (E), and stomatal conductance to water vapor (gs) of carambola trees grown in containers in a Krome very gravelly loam soil. In the orchard, SWD never reached a point where there was a decrease on ΨS, A, E, or gs. For carambola trees in containers, when SWD levels were above 50% there was a reduction in ΨS that subsequently reduced gs. A reduction in gs resulted in a linear decrease in E and a sharp decline in A when gs fell below 50 mmol•m-2•s-1. Leaf gas exchange was better correlated with ΨS than with SWD level. Therefore, ΨS may be a more accurate than SWD (as determined from measurements of soil water content) for irrigation scheduling of carambola in gravelly soils.
In the orchard, precipitation and possibly capillarity from the shallow water table resulted in sufficient soil water content to obtain adequate vegetative growth and yields. Adequate soil water content also resulted in no significant phenological differences among treatments. In containers where lateral water movement and capillary rise were prevented, tree trunk diameter and total dry weight were lower for trees irrigated at 32-50% SWD and 51-60% SWD than for trees irrigated at 0-21% SWD, 22-31% SWD treatments.
Based on the results of this study, irrigation of carambola trees in Krome soil at 17% SWD in an orchard and at 32% SWD in containers does not reduce tree growth, yield and fruit quality.
Carambola (Averrhoa carambola) is a commercially important tropical fruit tree in South Florida. Little is known about carambola water requirements or its response to limited soil water supply under subtropical conditions such as those of southern Florida. The objective of this study was to determine the effects of four levels of soil water depletion (SWD) that ranged from field capacity (FC) to the first visual signs of water stress (leaf yellowing and abscission), on leaf gas exchange, stem water potential, phenology, growth, and yield of 2-year-old, container-grown carambola trees, and 8-year-old 'Arkin' carambola trees in an orchard.
Soil water depletion treatments were determined from continuous measurement of soil water content using multisensor capacitance probes (EnviroSCAN, Sentek PTY Ltd., Kent Town, Australia). Irrigation of orchard trees was initiated when SWD reached one of the following levels (where 0% SWD = FC): 0-8% SWD, 9-11% SWD, 12-14% SWD, or 15-17% SWD. Trees in 95 L containers filled with Krome very gravelly loam soil were irrigated when SWD reached one of the following levels: 0-21% SWD, 22-31% SWD, 32-50% SWD, or 51-60% SWD.
An increase in SWD resulted in a reduction in stem water potential (ΨS), net CO2 assimilation (A), transpiration (E), and stomatal conductance to water vapor (gs) of carambola trees grown in containers in a Krome very gravelly loam soil. In the orchard, SWD never reached a point where there was a decrease on ΨS, A, E, or gs. For carambola trees in containers, when SWD levels were above 50% there was a reduction in ΨS that subsequently reduced gs. A reduction in gs resulted in a linear decrease in E and a sharp decline in A when gs fell below 50 mmol•m-2•s-1. Leaf gas exchange was better correlated with ΨS than with SWD level. Therefore, ΨS may be a more accurate than SWD (as determined from measurements of soil water content) for irrigation scheduling of carambola in gravelly soils.
In the orchard, precipitation and possibly capillarity from the shallow water table resulted in sufficient soil water content to obtain adequate vegetative growth and yields. Adequate soil water content also resulted in no significant phenological differences among treatments. In containers where lateral water movement and capillary rise were prevented, tree trunk diameter and total dry weight were lower for trees irrigated at 32-50% SWD and 51-60% SWD than for trees irrigated at 0-21% SWD, 22-31% SWD treatments.
Based on the results of this study, irrigation of carambola trees in Krome soil at 17% SWD in an orchard and at 32% SWD in containers does not reduce tree growth, yield and fruit quality.

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SOIL WATER DEPLETION, GROWTH, PHYSIOLOGY, AND YIELD OF CARAMBOLA TREES IN KROME SOIL

By RASHID AL-YAHYAI

A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY UNIVERSITY OF FLORIDA 2004

Copyright 2004 by Rashid Al-Yahyai

ACKNOWLEDGMENTS I would like to acknowledge the help and support of many individuals without whom this work would not have been possible. My sincere gratitude goes to my major professors, Drs. Bruce Schaffer and Frederick S. Davies, for their guidance and direction of matters related to my academic performance and professional development. I deeply appreciate the help and support of members of my supervisory committee Drs. Jonathan H. Crane, Yuncong Li, and Thomas Obreza. I also thank the following individuals for their assistance with the research project: Angel Colls, Osvany Rodriguez, Mark Kohout, and Dr. Rafael Muñoz-Carpena. I am thankful to many friends who made my stay in Florida most enjoyable.

TABLE OF CONTENTS page ACKNOWLEDGMENTS ................................................................................................. iv LIST OF TABLES............................................................................................................ vii LIST OF FIGURES ........................................................................................................... ix ABSTRACT..................................................................................................................... xiii 1 2 INTRODUCTION ........................................................................................................1 REVIEW OF LITERATURE .......................................................................................8 Carambola.....................................................................................................................8 Origin and Distribution..........................................................................................8 Economic Importance............................................................................................9 Botany and Uses ..................................................................................................10 Tree Phenology....................................................................................................14 Cultivation ...........................................................................................................15 Environmental Stress Physiology of Carambola.................................................23 Temperature .................................................................................................23 Wind .............................................................................................................25 Flooding .......................................................................................................26 Drought and water stress ..............................................................................26 Water Stress ................................................................................................................29 Effect of Water Stress on Vegetative Growth .....................................................32 Root growth and distribution........................................................................32 Leaf area .......................................................................................................32 Trunk growth................................................................................................33 Shoot growth ................................................................................................34 Effect of Water Stress on Reproductive Growth and Yield ................................34 Flowering and fruiting..................................................................................34 Yield .............................................................................................................36 Fruit quality ..................................................................................................36 Effect of Water Stress on Physiological Processes .............................................37 Net CO2 assimilation and transpiration........................................................37 Stomatal conductance...................................................................................38 Leaf and stem water potential ......................................................................39 Irrigation Scheduling of Fruit Trees ...........................................................................42 v

Irrigation scheduling based on plant water status ........................................43 Irrigation scheduling based on soil water status...........................................45 Conclusions.................................................................................................................50 3 CALIBRATION OF THE NEUTRON PROBE AND MULTISENSOR CAPACITANCE PROBES IN A GRAVELLY LOAM CALCAREOUS SOIL ......53

��� Materials and Methods ...............................................................................................55 ��� Results and Discussion ...............................................................................................59 ��� Conclusions .................................................................. ................................ .............61 ��� MONITORING SOIL WATER CONTENT FOR IRRIGATION SCHEDULING IN 4 ��� A CARAMBOLA ORCHARD IN A GRAVELLY LIMESTONE SOIL .................68

������������������������ ������ and Methods ...............................................................................................71 Materials ������������������������� ����� and Discussion ...............................................................................................73 Results

���������������������������������� 5 �� HYSIOLOGICAL RESPONSES OF CARAMBOLA TREES TO WATER ������������������������������������ DEPLETION IN KROME VERY GRAVELLY LOAM SOIL.................................83 ���� ������������������������������������������ � Materials and Methods ............................................................................... ................85 ������������������������������������������...........88 � Results and Discussion .................................................................................... ������������������������������������������ ..........91 � Conclusions.......................................................................................................

�������������������������������� ��������� � 6 GROWTH, YIELD, AND FRUIT QUALITY OF CARAMBOLA AS AFFECTED ���������������������������������������� ��� SOIL WATER DEPLETION................................................................... ..........101 BY

�������������������������������������� �������� and Methods ............................................................................. ................102 Materials ������������������������������������� ��������� Discussion ........................................................................... ..................105 Results and ������������������������������������� ��������� Conclusions............................................................................................ ...................112 ��������������������������������� ��������������������������������� ������ ���� HENOLOGICAL CYCLES OF CARAMBOLA AT FOUR LEVELS OF SOIL 7�� ���������������������������������������� ����� DEPLETION IN SOUTH FLORIDA ...................................... .................127 ATER

������������������������������������ ������������Methods .............................................................................................130 Materials and ������������������������������������ ������������ Results.......................................................................................................................133 ������������������������������������ ������������ Discussion.................................................................................................................137 ����������������������������������� ������������ onclusions.................................................................................................. ............141

���������������������������������� ������������� AND CONCLUSIONS .......................................................................149 8 SUMMARY

����������������������������������� ����������� SOIL WATER DEPLETION, PHYSIOLOGY, GROWTH, AND YIELD, � APPENDIX � �������� ���������������������������������� ������������CARAMBOLA TREES IN KROME SOIL...............� OF ......................152

������ ����������������������������� ������������������ LIST OF REFERENCES................................................................................................158

����������������������������������� ������������� SKETCH ..........................................................................................173 BIOGRAPHICAL vi

LIST OF TABLES Table page

2-1. Major carambola cultivars of economic importance in several major carambola production regions....................................................................................................21 3-1. Fitted parameters of van Genuchten’s model used to describe soil water retention curves of Krome very gravelly loam soil where soil water content was measured in a carambola orchard using multisensor capacitance probes (EnviroSCAN) at soil depths of 10, 20, and 30 cm. ....................................................................................62 3-2. Fitted parameters of van Genuchten’s model used to describe soil water retention curves of Krome very gravelly loam soil where soil water content was measured in a carambola orchard using a neutron probe at soil depths of 10, 20, and 30 cm. ....63 4-1. Fitted parameters of van Genuchten’s model used to create a soil water retention curves for Krome soil. Soil water content was measured in a carambola orchard with a neutron probe and multisensor capacitance probes. ......................................78 6-1. Effect of soil water depletion (SWD) on fresh and dry weights of leaves, stem, and roots of carambola trees in containers....................................................................114 6-2. Effect of soil water depletion (SWD) on total fruit number and fruit weight of carambola trees in an orchard.................................................................................115 6-3. Effect of soil water depletion (SWD) on fresh weight of mature fruit (108 ohue) of carambola trees in an orchard.................................................................................116 6-4. Effect of soil water depletion (SWD) on dry weight of mature fruit (108 ohue) of carambola trees in an orchard.................................................................................117 6-5. Effect of soil water depletion (SWD) on fresh weight of ripe fruit (tree-ripened, 87 o hue) of carambola trees in an orchard...................................................................118 6-6. Effect of soil water depletion (SWD) on dry weight of ripe fruit (tree-ripened, 87 o hue) of carambola trees in an orchard...................................................................119 6-7. Effect of soil water depletion (SWD) on fresh and dry weights of mature fruit (108 o hue) of carambola trees in containers. ..................................................................120

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6-8. Effect of soil water depletion (SWD) on fresh and dry weights of ripe fruit (treeripened, 87 ohue) of carambola trees in containers.................................................121 6-9. Effect of soil water depletion (SWD) on total soluble solid (TSS) content of ripe fruit (tree-ripened, 87 ohue) of carambola trees in an orchard........................................122 6-10. Effect of soil water depletion (SWD) on total soluble solid (TSS) content of ripe fruit (tree-ripened, 87 ohue) of carambola trees in containers. ...............................123 7-1. The effect of soil water depletion (SWD) on shoot flushing, axillary shoot growth, flowering, and fruit load of orchard-grown carambola trees in 2003. ...................142 7-2. The effect of soil water depletion (SWD) on shoot flushing, axillary shoot growth, and flowering of container-grown carambola trees in 2003...................................143 A-1. Effect of soil water depletion (SWD) on total number and weight of carambola trees grown in containers. ...............................................................................................152 A-2. Average soil water depletion (SWD) prior to irrigation of carambola trees in containers in the field and in an orchard in 2002 and 2003. ..................................154

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LIST OF FIGURES Figure page

3-1. Relationship between scaled frequency (SF) from a multisensor capacitance probe (EnviroSCAN) and volumetric water content (%) fitted to obtain calibration coefficients for Krome very gravelly loam soil in containers, y = 0.011x + 0.5206; r2 = 0.98. ...................................................................................................................64 3-2. Relationship between count ratio from the neutron probe and volumetric water content (%) fitted to obtain a calibration equation for Krome very gravelly loam soil in containers, y = 29.05x - 6.4; r2 = 0.95. ..........................................................65 3-3. Soil water retention curves for Krome very gravelly loam soil at 10-, 20-, and 30-cm depths based on van Genuchten’s model calculated from volumetric water content (θv) measured with capacitance probes and soil water tension measured with tensiometers..............................................................................................................66 3-4. Soil water retention curves for Krome very gravelly loam soil at 10-, 20-, and 30-cm depths based on van Genuchten’s model calculated from volumetric water content (θv) measured with a neutron probe and soil water tension measured with tensiometers..............................................................................................................67 4-1. Total monthly rainfall and evapotranspiration (ET) during 2001 and 2002 in Homestead, Fla. Source: Florida Automated Weather Network, IFAS, Univ. of Fla., Gainesville................................................................................................................79 4-2. Relationship between volumetric soil water content measured with a neutron probe and soil water tension measured with a tensiometer in a carambola orchard in Krome very gravelly loam soil. The response line was fitted using the van Genuchten equation (r2 = 0.42). ...............................................................................80 4-3. Relationship between volumetric soil water content measured with multisensor capacitance probes and soil water tension measured with a tensiometer in a carambola orchard in Krome very gravelly loam soil. The response line was fitted using the van Genuchten equation (r2 = 0.35)..........................................................81 4-4. Relationship between soil water content (%) as measured by multisensor capacitance probes and a neutron probe in Krome very gravelly loam soil (y = 0.67 x + 12.19, r2 = 0.35). ................................................................................82

ix

5-1. Soil water depletion (SWD) and stem water potential (Ψs) of carambola trees in containers in a greenhouse (y = -0.44 - (3.59e-06) x3, r2 = 0.82). ............................93 5-2. Soil water depletion (SWD) and stem water potential (Ψs) of carambola trees in containers in the field (y = -0.48 - (2.79e-06) x3, r2 = 0.56).....................................93 5-3. Soil water depletion (SWD) and stomatal conductance to water vapor (gs) of carambola trees in containers in a greenhouse (y = 95.46 – 0.0002 x3, r2 = 0.46). ..94 5-4. Stem water potential (Ψs) and stomatal conductance to water vapor (gs) of carambola trees in containers in a greenhouse (y = 11.88 + 132.94 ex , r2 = 0.54)....................95 5-5. Stem water potential (Ψs) and stomatal conductance to water vapor (gs) of carambola trees in containers in the field (y = 44.76 – 41.57 / x, r2 = 0.32)..............................95 5-6. Stem water potential (Ψs) and transpiration (E) of carambola trees in containers in a greenhouse (y = 0.30 + 2.94 ex, r2 = 0.55)................................................................96 5-7. Stem water potential (Ψs) and transpiration (E) of carambola trees in containers in the field (y = 1.03 + 2.11 ex, r2 = 0.28).....................................................................96 5-8. Stomatal conductance to water vapor (gs) and transpiration (E) of carambola trees in containers in a greenhouse (y = 0.33 + 0.02 x, r2 = 0.76).........................................97 5-9. Stomatal conductance to water vapor (gs) and transpiration (E) of carambola trees in containers in the field (y = 0.34 + 0.02 x, r2 = 0.75). ...............................................97 5-10. Stem water potential (Ψs) and net CO2 assimilation (A) of carambola trees in containers in a greenhouse (y = -0.09 + 9.76 ex, r2 = 0.70). .....................................98 5-11. Stem water potential (Ψs) and net CO2 assimilation (A) of carambola trees in containers in the field (y = 1.19 - 2.95/x, r2 = 0.31). ................................................98 5-12. Stomatal conductance (gs) and net CO2 assimilation (A) of carambola trees in containers in a greenhouse (y = -2.46 + 0.39 (lnx)2, r2 = 0.75). ...............................99 5-13. Stomatal conductance (gs) and net CO2 assimilation (A) of carambola trees in containers in the field (y = -2.24 + 0.40 (lnx)2, r2 = 0.45). .......................................99 5-14. Leaf (ΨL) and stem water potentials (ΨS) of carambola trees in containers in the field (y = 0.10 + 1.00x, r2 = 0.97) and in a greenhouse (y = 0.33 + 0.91x, r2= 0.97)....................................................................................100 5-15. Leaf (ΨL) and stem water potentials (ΨS) of carambola trees in an orchard in the field (y = 0.73 + 1.38x, r2 = 0.85)...........................................................................100

x

6-1. Effect of soil water depletion (SWD) on shoot length of orchard-grown carambola trees during 2002 (A) and 2003 (B). Symbols with vertical bars represent means ± SE of 4 shoots per tree for 6 trees. .........................................................................124 6-2. Effect of soil water depletion (SWD) on trunk diameter of young carambola trees grown in containers. Symbols with vertical bars represent means ± SE of 6 trees. ................................................................................................................125 6-3. Effect of soil water depletion (SWD) on fruit length of orchard-grown carambola trees during 2003 summer harvest (August) and winter harvest (December). Symbols with vertical bars represent means ± SE of 4 fruit per tree for 6 trees....126 7-1. Average 41-year and 2003 monthly precipitation and average 2003 monthly temperature in Homestead, Fla. Source: Florida Automatic Weather Network (FAWN) (2003) and The National Oceanic and Atmospheric Administration (NOAA) (2003). .....................................................................................................144 7-2. Shoot flush ratings of carambola trees in Krome very gravelly loam soil of South Florida in 2003 irrigated at four levels of soil water depletion (SWD) in an orchard (A, n = 51) and in containers (B, n = 22). ..............................................................145 7-3. Shoot growth ratings of carambola trees in Krome very gravelly loam soil of South Florida in 2003 irrigated at four levels of soil water depletion (SWD) in an orchard (A, n = 51) and in containers (B, n = 22). ..............................................................146 7-4. Flowering ratings of carambola trees in Krome very gravelly loam soil of South Florida in 2003 irrigated at four levels of soil water depletion (SWD) in an orchard (A, n = 51) and in containers (B, n = 22). ..............................................................147 7-5. Fruit load rating of carambola trees in Krome very gravelly loam soil of South Florida in 2003 irrigated at four levels of soil water depletion (SWD) in an orchard (A, n = 51) and in containers (B, n = 22). ..............................................................148 A-1. Soil water depletion treatments (SWD) as determined from measurements of soil water content using multisensor capacitance probes (EnviroSCAN) in an 8-year-old carambola orchard in Krome soil in South Florida. Field capacity was determined based on the pattern of soil water depletion over a period of 7 d. Visual stress symptoms (VSS) occurred when the leaves became wilted and chlorotic.............153 A-2. Vapor pressure (VP) difference between saturated and ambient air vapor pressure in the greenhouse and the orchard..............................................................................155 A-3. Soil water depletion (SWD) as measured with capacitance probes (EnviroSCAN) in an 8-year-old carambola orchard in Krome soils. Points of ‘field capacity’ and ‘onset of water stress’ were determined based on the pattern of soil water depletion. Measurements of stem water potential (ΨS) and net CO2 assimilation (A) indicated by the vertical double-line on the x-axis were chosen randomly at a point beyond the theoretical ‘onset of water stress’ point............................................................156 xi

A-4. Ratings of the phenological cycles of carambola trees in Krome very gravelly loam soil in an orchard in South Florida in 2003 Trees were irrigated at four levels of soil water depletion (SWD) in an orchard (n = 51).......................................................157

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Abstract of Dissertation Presented to the Graduate School of the University of Florida in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy SOIL WATER DEPLETION, GROWTH, PHYSIOLOGY, AND YIELD OF CARAMBOLA TREES IN KROME SOIL By Rashid Al-Yahyai December 2004 Chair: Bruce Schaffer Cochair: Frederick S. Davies Major Department: Horticultural Science Carambola (Averrhoa carambola) is a commercially important tropical fruit tree in South Florida. Little is known about carambola water requirements or its response to limited soil water supply under subtropical conditions such as those of southern Florida. The objective of this study was to determine the effects of four levels of soil water depletion (SWD) that ranged from field capacity (FC) to the first visual signs of water stress (leaf yellowing and abscission), on leaf gas exchange, stem water potential, phenology, growth, and yield of 2-year-old, container-grown carambola trees, and 8-yearold 'Arkin' carambola trees in an orchard. Soil water depletion treatments were determined from continuous measurement of soil water content using multisensor capacitance probes (EnviroSCAN, Sentek PTY Ltd., Kent Town, Australia). Irrigation of orchard trees was initiated when SWD reached one of the following levels (where 0% SWD = FC): 0-8% SWD, 9-11% SWD, 12-14% SWD, or 15-17% SWD. Trees in 95 L containers filled with Krome very gravelly loam soil xiii

were irrigated when SWD reached one of the following levels: 0-21% SWD, 22-31% SWD, 32-50% SWD, or 51-60% SWD. An increase in SWD resulted in a reduction in stem water potential (ΨS), net CO2 assimilation (A), transpiration (E), and stomatal conductance to water vapor (gs) of carambola trees grown in containers in a Krome very gravelly loam soil. In the orchard, SWD never reached a point where there was a decrease on ΨS, A, E, or gs. For carambola trees in containers, when SWD levels were above 50% there was a reduction in ΨS that subsequently reduced gs. A reduction in gs resulted in a linear decrease in E and a sharp decline in A when gs fell below 50 mmol·m-2·s-1. Leaf gas exchange was better correlated with ΨS than with SWD level. Therefore, ΨS may be a more accurate than SWD (as determined from measurements of soil water content) for irrigation scheduling of carambola in gravelly soils. In the orchard, precipitation and possibly capillarity from the shallow water table resulted in sufficient soil water content to obtain adequate vegetative growth and yields. Adequate soil water content also resulted in no significant phenological differences among treatments. In containers where lateral water movement and capillary rise were prevented, tree trunk diameter and total dry weight were lower for trees irrigated at 3250% SWD and 51-60% SWD than for trees irrigated at 0-21% SWD, 22-31% SWD treatments. Based on the results of this study, irrigation of carambola trees in Krome soil at 17% SWD in an orchard and at 32% SWD in containers does not reduce tree growth, yield and fruit quality.

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CHAPTER 1 INTRODUCTION Carambola (Averrhoa carambola L.), also called star fruit, is a member of the Oxalidaceae and a native of Southeast Asia. This tropical fruit tree is cultivated between 30o latitude (Watson et al., 1988), primarily in Southeast Asia and also in China, India (Nand, 1970; Popenoe, 1948; Tidbury, 1976), Guyana, Australia, Israel, and the United States (mainly Hawaii and Florida) (Crane, 1994). There are approximately 100 ha of carambola trees in Florida (J. H. Crane, personal communication), of which 46 ha are in Miami-Dade County (Degner et al., 2002). A sweet-type, ‘Arkin’, has been the leading commercial carambola cultivar in Florida (Campbell, 1989; Knight and Crane, 2002; Lamberts and Crane, 1990; Núñez-Elisea and Crane, 1998). Carambola trees are adapted to various types of well-drained soils (Crane, 1994); however, deep, rich, and fertile soils of loamy-sand or clay-loam texture are generally favorable for optimum growth and production (Galán Saúco et al., 1993). Environmental factors such as low temperature (Campbell and Malo, 1981; Campbell et al. 1985; George et al., 2002a, 2002b; Watson et al., 1988), drought (Ismail and Awang, 1992; Ismail and Noor, 1996a; Ismail et al., 1994, 1996; Salakpetch et al., 1990), flooding (Ismail and Noor, 1996b; Joyner and Schaffer, 1989), sunlight, (Marler et al., 1994) and wind (Campbell, 1985; Crane, 1993; Crane et al., 1994a; Marler and Zozor, 1992; Morton, 1987), as well as cultural practices such as cultivar selection (Salakpetch et al., 1990), mulching (George et al., 2000, 2001), fertilization (Campbell, 1985, 1989; Crane, 1994; Knight, 1982), and pruning (Núñez-Elisea and Crane, 1998, 1

2 2000) influence vegetative and reproductive growth of carambola trees. In their native tropical habitat, trees grow and bear fruit year-round (Núñez-Elisea and Crane, 2000). Trees may bear new flushes, flowers, and immature and mature fruit simultaneously (Galán Saúco et al., 1993). However, this phenological cycle can be interrupted due to unfavorable environmental conditions during the winter months that include low temperatures, dry winds, and high light intensity (Núñez-Elisea and Crane, 1998). Tree responses to water stress depend on the stage of growth and duration and severity of the stress. Vegetative growth of carambola was reduced with increased water stress (Ismail and Awang, 1992; Ismail et al., 1994, 1996). The response of carambola tree flowering to water stress depends on the time of stress. Flowering of carambola is more affected by water stress and cultivar than by diurnal temperature variation and photoperiod (Salakpetch et al., 1990). Even when tree growth is not affected by drought, yield was significantly reduced in rain-fed trees compared to irrigated trees, mainly due to a decrease in fruit size (Bookeri, 1996). Leaf water potential (ΨL) decreases when water loss via transpiration (E) is greater than water uptake (Salisbury and Ross, 1985). Studies on carambola have shown that ΨL declines with increasing water stress (Ismail and Noor, 1996b; Ismail et al., 1994, 1996). Salakpetch et al. (1990) observed that leaves of stressed carambola trees wilted when midday ΨL reached -2.0 MPa and became yellow and abscised 5 d later. However, trees appeared normal and rapidly recovered to their original ΨL levels of -0.6 MPa 2 to 3 d after rewatering. Ismail et al. (1994) found that net CO2 assimilation (A) of B17 carambola clones grafted onto B2 clones in a greenhouse declined as a result of water

3 stress. Stomatal conductance (gs) was two to three times lower in stressed trees compared to control trees. Irrigation scheduling involves regulating the amount and frequency of irrigation to meet the water demand of a particular crop. Application of water may vary depending on plant needs which may vary depending on growth stage, climatic conditions, and soil type. Regulation of irrigation is necessary to avoid over-watering fruit orchards that may result in reduced tree growth and production from waterlogging, loss of water through runoff or deep percolation, waste of energy resources, and potential agrochemical leaching. Efficient irrigation management helps reduce the effects of insufficient water supply that results in plant stress and a subsequent reduction in growth, yield and fruit quality. The amount of water needed depends mainly on soil evaporation and plant transpiration (evapotranspiration, ET). Drought (Ismail and Noor, 1996a; Ismail et al., 1996; Salakpetch et al., 1990) and excessive soil water (Ismail and Noor, 1996b; Joyner and Schaffer, 1989) reduce carambola growth and yield. In some areas, scheduling irrigation is vital for commercial carambola production. However, little is known about irrigation scheduling practices in carambola producing countries. In south Florida, Crane (1994) recommended applying 33 mm of water per ha twice a week during dry periods throughout the year. However, irrigation rates and frequencies needed for optimum growth and yield have not been established. Most of the annual rainfall in Florida occurs during the summer months. Historical climatic records for south Florida show that about 80% of rainfall occurred between May and October in 2001 to 2003. During winter, irrigation is used to compensate for the low rainfall and for

4 freeze protection. Irrigation is also used to compensate for unevenly distributed rainfall within a month. A survey by Li et al. (2000) in 1998 showed that 73% of tropical fruit growers of Miami-Dade County schedule irrigation based on the frequency and quantity of rain. The percentage declined to 64.3% in 2002 according to a more recent survey by Muñoz-Carpena et al. (2003). The number of respondents who monitored soil water content for irrigation scheduling increased to 48.8% in 2002 (Muñoz-Carpena et al., 2003) compared to only 15% in 1998 (Li et al., 2000). Problems concerning variability in irrigation duration and frequency were rated high among tropical fruit growers (Li et al., 2000), which highlights the need for a better understanding of irrigation requirements of these crops. Tensiometers, neutron probes, and capacitance probes directly or indirectly monitor soil water content and are often used for irrigation scheduling. Tensiometers measure soil suction or matric water potential rather than soil water content (Richards, 1942; Smajstrla and Harrison, 1998). A soil water retention curve must be established to determine the soil water content that corresponds to the water matric potential in order to estimate soil water content with a tensiometer. The neutron probe is a portable compact unit that is easy to operate, and volumetric soil water content can be obtained instantly at different depths in the soil profile. Neutron probes are considered more accurate than many other soil water status monitoring devices for irrigation scheduling (Evett and Steiner, 1995; Mostert and Hoffman, 1996). A “count rate” obtained from a neutron probe inserted into access tubes in the field is converted to volumetric water content using a calibration curve. Neutron probes measure soil volumetric water content as a percentage of water per volume of soil. A neutron probe contains radioactive materials which can cause health

5 and regulatory concerns; it is also relatively costly to purchase and maintain. Capacitance probes measure the soil water content based on the dielectric constant of the soil (Paltineanu and Starr, 1997; Phene et al., 1990; Wu, 1998), which includes dielectric constants of water (80.4), soil particles (3 to 7) and air (1) (Robinson and Dean, 1993; Paltineanu and Starr, 1997; Starr and Paltineanu, 1998a, 1998b; Wu, 1998). Since the dielectric constant of the soil particles and air are small and relatively constant compared to that of water, changes in the dielectric constant of the soil are a measure of the change in soil water content. The volumetric water content can be expressed either as a percentage at specific depths or total amount of water per volume of soil (Núñez-Elisea et al., 2001; Paltineanu and Starr, 1997; Starr and Paltineanu, 1998a). An advantage of capacitance probes system for measuring soil water content is that it provides continuous monitoring. However, initial capital investment and maintenance are costly for this system. Soil water content should be correlated with physiological responses, growth, and fruit production of tropical fruit crops, such as carambola, to determine the appropriate amount of water to apply to the crop. The response of fruit trees to soil water deficit is different from that of annual herbaceous plants. Trees are able to tolerate longer periods of low soil water content through specialized short-term and long-term physiological, phenological, anatomical, and morphological responses (Ludlow, 1989). Growth reduction, stomatal closure, and osmotic adjustment are among the physiological responses to a lower water potential (Hsiao, 1973; Turner, 1979). The correlation of soil water depletion to tree physiological processes, growth and fruit production provides a tool for accurate determination of tree water requirements. This aids in developing

6 irrigation scheduling strategies such as regulated deficit irrigation to obtain optimum yield and enhance fruit quality. Regulated deficit irrigation has been used for several fruit crops including peach (Chalmers et al., 1984), citrus (Castel and Buj, 1990; Domingo and Ruiz-Sanchez, 1996), and apple (Mills et al., 1997). Carambola trees respond to water deficit by drought avoidance (Neuhaus, 2003). This strategy involves minimizing water loss by stomatal control, leaf movement, and reduced leaf area (Ludlow, 1989). The lethal level of ΨL in carambola occurs at about -2.9 MPa and the lethal leaf relative water content (RWC) at about 56% (Ismail et al., 1994). The response of carambola trees to soil water deficit can be divided into four phases (Neuhaus, 2003): (1) reduction in osmotic adjustment and low osmotic adjustment; (2) increased root:shoot ratio by shedding leaves and fruit and increased leaf osmotic adjustment; (3) irreversible damage caused by chlorophyll degradation, root growth inhibition, and further leaf and fruit shedding; and (4) attainment of the lethal physiological state is reached. Rewatering the trees at the second phase results in the recovery of the plant to the original physiological state in a relatively short time (Salakpetch et al., 1990). Thus, correlating physiological changes to soil water depletion is essential to optimize irrigation amount and frequency. The objective of this research project was to relate soil water contents in a gravelly loam soil to physiology, growth, and yield of carambola trees. Studies were designed to determine the effects of different soil water depletion levels on physiology, growth and yield of young, container-grown, and mature, field-grown 'Arkin' carambola trees. The main hypothesis was that physiology, growth, and yield responses of carambola trees are closely related to soil water content.

7 Specific project hypotheses and questions addressed in this dissertation include the following: Hypothesis 1: Soil moisture measurements using multisensor capacitance probes and a neutron probe are site specific. The following question is addressed in chapter 3: What are the calibration equations for a neutron probe and multisensor capacitance probes (EnviroScan) in Krome very gravelly loam soil? Hypothesis 2: Tensiometers, multisensor capacitance probes, and neutron probes differ in their accuracy and effectiveness for assessing soil water content. The following question is addressed in chapter 4: What is the most accurate and practical technique for determining soil water content in a Krome very gravelly loam soil? Hypothesis 3: Water stress due to low soil water availability decreases leaf gas exchange and water potential of carambola trees. The following question is addressed in chapter 5: What is the effect of soil water depletion on A, gs, E and ΨS of carambola trees? Hypothesis 4: Reduced soil water availability reduces carambola growth, yield, and fruit quality. The following question is addressed in chapter 6: What is the effect of four soil water depletion levels on carambola vegetative growth, fruit growth, yield, and fruit quality? Hypothesis 5: Reduced soil water availability affects carambola phenology. The following question is addressed in chapter 7: What is the effect of four soil water depletion levels on carambola tree phenology?

CHAPTER 2 REVIEW OF LITERATURE Carambola Origin and Distribution Carambola (Averrhoa carambola L.) is one of two species of the two genera of woody plants in the Oxalidaceae, a family composed of primarily herbaceous species. Another woody species in the same genus, Averrhoa bilimbi L., also produces edible fruit commonly called bilimbi (Nakasone and Paull, 1998). The exact origin of carambola is not known. It is believed to be native to southeastern Asia and probably originated in Indochina or Indonesia (Galán Saúco et al., 1993; George and Nissen, 1994), or Sri Lanka and Moluccas (Crane, 1994 and Morton, 1987). Carambola is cultivated within 30o north and south of the equator (Watson et al., 1988; Galàn Saúco et al., 1993), primarily in Southeast Asia, Indonesia, Malaysia, and Thailand, but also in China, Taiwan, India (Nand, 1970; Popenoe, 1948; Tidbury, 1976), Brazil, Mexico, Guyana, Australia, Israel, and the United States (mainly Hawaii and Florida) (Crane, 1994; Mauro Pace, 2004, personal communication). Carambola grows best in tropical lowland areas with uniform medium to high rainfall throughout the year (Campbell and Malo, 1981; Martin et al., 1987; Popenoe, 1948; Samson, 1980; Tidbury, 1976). However, it can also grow well in humid, subtropical areas (Campbell and Malo, 1981) as well as in areas with seasonal periods of drought (Galán Saúco et al., 1993; Martin et al., 1987; Tidbury, 1976).

8

9 Economic Importance The spread of carambola as a commercial tropical fruit tree was due to the improvement in postharvest and cultivation practices, such as the control of fruit fly and the selection of sweet cultivars with high productivity and good quality (Galàn Saúco et al., 1993). No collective data on worldwide carambola production exist, but estimates from individual countries have been reported. According to Galàn Saúco et al. (1993) and Mauro Pace (2004, personal communication), major carambola producing countries are Indonesia, Malaysia, Taiwan, Thailand; emerging producers are Brazil, Mexico and, more generally, Caribbean countries. The current estimate of world production is 150,000 to 200,000 metric tons (MT) (Mauro Pace, 2004, personal communication). Carambola production in Indonesia, which includes wild trees, fluctuated from 50,000 to 55,000 MT between 1995 and 2000. Commercial production in Malaysia was 8,600 MT in 20002002, declining from 29,000 MT in 1990-91. Malaysian production estimates, however, may differ according to sources, in some cases reaching 40,000 MT, including production from wild trees and non-commercial production. In Taiwan, production was estimated to be 25,000 MT on 1,000 ha in 2002, declining from an area of 3,100 ha with 38,270 MT produced in 1986 (Galàn Saúco et al., 1993). In 2001, production in Thailand was estimated to be about 20,000 MT, Peru produced 1,766 MT, Israel produced 200 MT, and commercial production from the USA (Florida and Hawaii) was estimated at about 4,000 MT (Mauro Pace, 2004, personal communication). In the United States, the greatest hectarage of carambola occurs in Florida with an estimated 100 ha in 2002 (Jonathan H. Crane, 2003, Personal communication) with estimated annual production of 36.5 MT per ha. Hawaii has an estimated 8 ha of carambola with an annual production of 43.5 MT per ha (H.A.S.S., 2001). Carambola

10 production in south Florida increased in the late 1980s and early 1990s as improved cultivars such as ‘Arkin’ and aggressive consumer marketing created interest among growers (Degner et al., 2002; Knight et al., 1984). In addition, an increase in immigration from Southeast Asia and the recognition by growers and packers of the productivity and market potential of this fruit led to increased planting of carambola (Knight et al., 1984). By 1990, an estimated 242 ha had been planted in Miami-Dade County in Florida. In the years following Hurricane Andrew in 1992, hectarage in Miami-Dade County began to decline until there were only about 46 ha in 2001 (Degner et al., 2002). Despite the decline in hectarage, carambola continued to be one of the highest value crops in south Florida making it an economically viable fruit crop (Degner et al., 2002). Botany and Uses Growth habit. Carambola is a short-trunk evergreen tree with a broad, rounded canopy (Morton, 1987; and Crane, 1994; Nakasone and Paull, 1998). Branches of carambola are plagiotropic and tree height increases by continual superposition of major limbs one on top of the other in a growth pattern that conforms to Troll’s model (Crane et al., 1991; Crane et al., 1994a). According to Troll’s growth model, non-trained trees have multiple leaders with an initial vertical orientation. Upon fruiting, the upright branches become laterally oriented and new shoots develop along these laterally oriented limbs (Halle et al., 1978). This results in overlapping branches and a spreading growth pattern with the interior canopy becoming defoliated (Crane et al., 1991; Crane et al., 1994a). Carambola trees possess a deep taproot and extensive lateral roots that develop a very dense mass of thin roots with profuse absorbent hairs (Galàn Saúco et al., 1993). Due to the gravelly texture of South Florida soils, which prevents deep penetration of the root system, roots in South Florida orchards are concentrated in the top 10 to 20 cm of the soil

11 profile (Núñez-Elisea et al., 2001). Due to the hardness of the soil in southern Florida, tropical trees including carambola are grown at intersections of perpendicular trenches that are 40 cm wide and 50 to 60 cm deep (Bünemann and Crane, 2000; Colburn and Goldweber, 1961; Núñez-Elisea et al., 2001). Most of the roots in Krome soil are confined to the trenches and no taproot is evident; instead, numerous short verticallyoriented roots grow to the bottom of the trench (Crane et al., 1994a). However, roots radiate laterally in all directions in the “plow” layer (10 cm deep). Root restriction can lead to reduced leaf and root growth especially when trees are water stressed (Ismail and Noor, 1996a). In Florida, vegetative and reproductive growth of carambola is vigorous from April to October. After the October harvest, trees bloom and vegetative growth slows down until it eventually ceases from November to February (Núñez-Elisea and Crane, 1998). During this period, leaves often exhibit interveinal chlorosis, which is a typical symptom of iron deficiency, and the rate of leaf senescence increases resulting in partial or full defoliation (Núñez-Elisea and Crane, 1998). Leaves. Leaves of carambola trees are compound, spirally arranged and alternate, 15 to 30 cm long, with 5-11 nearly opposite, ovate-oblong leaflets that are 1.5 to 9.0 cm long and 1.0 to 4.5 cm wide (Campbell et al., 1985; Galán Saúco et al., 1993). They are medium-green, and smooth on the upper surface, and slightly pubescent and whitish on the lower surface (Crane, 1994; Morton, 1987). Flowers. Carambola flowers are 5 to 11 mm long, 5 mm in diameter, with pink to lavender petals, perfect and borne in clusters in axils of leaves on young branches or on older branches without leaves. Most flowering and fruit production occur at the middle

12 canopy level (Crane et al., 1991). Carambola trees are heterostylous. Some cultivars bear flowers with long styles and short stamens, while others have short styles and long stamens (Campbell et al., 1985; Crane, 1994; Galán Saúco et al., 1993; Morton, 1987). In tropical areas, carambola trees have the potential to flower year round (Samson, 1980; Tidbury, 1976). Núñez-Elisea and Crane (1998) reported that pruning could stimulate year-round flowering. They found that buds older than 10 weeks produced panicles, while 6-week-old buds produced vegetative growth. Carambola flowers open in the morning between 0800 and 1000 HR and close in the afternoon between 1400 and 1800 HR the same day (Salakpetch et al., 1990). Due to their bright color, availability of nectar, and pollen viscosity, carambola flowers are attractive to bees (Apis spp.), which are essential for pollination. However, flowers may also be partially wind pollinated since some pollen is released outwards at the moment when anther dehiscence occurs (Galán Saúco et al., 1993). Some flowers may require cross-pollination to attain adequate fruit set and yield because of flower heterostyly (Knight, 1965). However, cross-pollination by cultivars with different stylar types is not always necessary in ‘Arkin’ and ‘Golden Star’ because they produce an abundant fruit crop when planted in solid blocks (Crane, 1994). Fruit. Carambola fruit are oval to ellipsoid, 5-15 cm long, and 3-10 cm in diameter, with 4-6 (usually 5) prominent longitudinal ribs. Fruit slices cut in cross-section are star-shaped, hence the name star fruit. The skin is thin, yellow to dark yellow and smooth with a waxy cuticle. Immature fruit are green, turning yellow or orange when ripe. The flesh is light-yellow to yellow, translucent, crisp, very juicy, and is not fibrous. Depending on the cultivar, the fruit vary from very sour with little sugar, to very sweet

13 with little acidity. It takes about 60 to 75 d from fruit set for the fruit to reach maturity (Campbell et al., 1985; Crane, 1994; Galán Saúco et al., 1993). Most of the fruit (>75%) is produced on the periphery of the tree; therefore, it is subject to ‘wind-scar’ from dead twigs under windy conditions (Núñez-Elisea and Crane, 1998, 2000). The potential annual fruit yield of carambola ranges from 4.5-18 kg/tree in the 2nd to 3rd years, 45 to 68 kg/tree in the 5th to 6th year, and 112-160 kg/tree at maturity between the 7th and 12th year (Campbell 1989; Crane, 1993,1994; Núñez-Elisea and Crane, 1998). Carambola seeds are small, 0.6 to 1.3 cm long, thin, brown, edible, and there are usually 10 to 12 seeds per fruit (Morton, 1987). In subtropical climates, some fruit have 1 to 2 seeds (Galán Saúco et al., 1993), while some fruit are seedless (Crane, 1994). Seeds are light brown and enclosed by a gelatinous aril. Seeds lose their viability in a few days after removal from the fruit (Crane, 1994). Uses. Carambola fruit from sweet cultivars are mainly consumed fresh; however, the fruit may be frozen, pickled, preserved, or used in various culinary preparations (Crane 1994; Cooper et al., 1998). Fruit of acidic cultivars are consumed cooked (Galàn Saúco et al., 1993). Most of Florida carambola fruit are marketed for fresh consumption. Approximately 20% to 40% of the fruit produced do not meet the fresh market standards, but can be utilized for processing (Matthews et al., 1988). Carambola fruit are a good source of potassium and a moderate source of vitamin C (Cooper et al., 1998). One hundred grams of the edible portion contains 35.7 calories, 8991g water, 0.38g protein, 0.08g fat, 9.38g carbohydrates, 0.8-0.9g fiber, 4.4-6mg calcium,

14 15.5-21mg phosphorus, 0.32-1.65mg iron, 26-53.1mg ascorbic acid, and trace amounts of carotene, thiamine, riboflavin, and niacin (Morton, 1987). Tree Phenology Cull (1986) divided the phenological cycle of tropical evergreen trees into three phases: the vegetative phase, the flower development phase, and the root development phase. Each phase has an effect on the other phases. For example, vegetative growth is essential for carbohydrate accumulation that is required for the development of young, photosynthetically-efficient leaves, root growth, vegetative growth, and fruit growth and development. Environmental factors such as low temperatures (Campbell and Malo, 1981; Campbell et al., 1985; Crane, 1994; George et al., 2002a, 2002b; Watson et al., 1988), drought (Ismail and Noor, 1996a; Ismail and Awang, 1992; Ismail et al., 1994, 1996; Salakpetch et al., 1990) , flooding (Ismail and Noor, 1996b; Joyner and Schaffer, 1989), sunlight, (Marler et al., 1994) and wind (Campbell, 1985; Crane, 1993; Crane et al., 1994a; Marler and Zozor, 1992; Morton, 1987), as well as cultural practices such as cultivar selection (Salakpetch et al., 1990), fertilization (Campbell, 1985,1989; Crane, 1994; Knight, 1982), mulching (George et al., 2000; 2001), and pruning (Núñez-Elisea and Crane, 1998, 2000), influence vegetative and reproductive growth of carambola. In their native tropical habitat, carambola trees grow and bear fruit year-round (Núñez-Elisea and Crane, 2000). Trees may produce new flushes, flowers, and bear immature and mature fruit simultaneously (Galán Saúco et al., 1993). Under southern Florida’s winter conditions that include low temperatures, dry winds, and high light intensities (above 1000 µmol·m-2·s-1), canopies become chlorotic and defoliate from February to March (Núñez-Elisea and Crane, 1998). Trees remain visually quiescent with

15 complete or partial defoliation during cool weather and begin to re-foliate from March to mid-May and may continue to produce new leaves to mid-October (Núñez-Elisea and Crane, 1998). Shoot growth alternates with periods of quiescence and apical growth results in an extension of the shoot axis. Each growth flush contains between eight and 12 leaves (Núñez-Elisea and Crane, 1998). Shoots that are formed during the spring have four to five flushes but those formed in the late summer normally flush once before shoot extension growth ceases in the fall (Núñez-Elisea and Crane, 1998). Carambola trees have several flowering periods and may flower continuously during the year under favorable cultural and environmental conditions (Núñez-Elisea and Crane, 1998, 2000). Two reproductive growth phases were observed in South Florida. The first phase is characterized by peaks of flowering in May and fruit maturity from July to October. The second phase is characterized by a peak bloom in September that results in fruit maturity from December to February (Campbell et al., 1985; Núñez-Elisea and Crane, 1998). In south Florida, the flowering phase of carambola extends from March to early November, and production season extends from mid-July to mid-February (Campbell et al., 1985; Núñez-Elisea and Crane, 1999, 2000). Núñez-Elisea and Crane (1998) concluded that growth promoting environmental conditions such as water availability or cultural practices such as pruning, defoliation, and nitrogen application impact floral initiation more directly than tree carbohydrate status. Cultivation Soil requirements. Carambola trees are adapted to various types of well-drained soils (Crane 1994; Popenoe, 1948; Nakasone and Paull, 1998). However, deep, rich, and fertile soils with loamy-sand or clay-loam texture are generally favorable for optimum growth and production (Galán Saúco et al., 1993). Soil pH between 5.5 and 6.5 is

16 generally ideal for tree growth (Campbell and Malo, 1981; Crane, 1994; Galán Saúco et al., 1993; Green, 1987). In acid soils in tropical climates, carambola trees grow vigorously and produce large deep-green leaves with few micronutrient deficiency symptoms. Symptoms of micronutrient deficiency are more prevalent in calcareous soils (Green, 1987). Carambola trees can tolerate soils with a pH below 5.5 much better than avocado, mango or lytchee (Galán Saúco et al., 1993). In fact, carambola trees are grown in pH 4.5 soils in Selangor State, Malaysia (Green, 1987) and in oolitic-limestone soils with a pH between 7.5 and 7.8 in south Florida (Knight, 1982). However, due to the high alkalinity of limestone soils, application of chelated iron, manganese, and zinc are necessary to prevent micronutrient deficiencies (Crane, 1994; Knight, 1982). In Miami-Dade County, Florida, carambola trees are generally grown in Krome very gravelly loam soils. This mineral soil is very shallow, with a pH of 7.4 to 8.4 (Noble et al., 1996). The soil is extremely low in organic matter and commercial farming largely depends on fertilizer applications (Degner et al., 2002). Krome soil is very porous with a gravelly loam surface layer between 8 and 23 cm thick, below which is a hard but porous layer of limestone bedrock (Degner et al., 1997; Degner et al,, 2002; Moseley et al., 1990). Planting density. Spacing varies among cultivars, soil types and pruning practices. A minimum distance of 5-7 m between trees and 6-9 m between rows is recommended (Nakasone and Paull, 1998). In South Florida, carambola trees are usually planted at moderate densities. Trees are planted at 4.6 to 6.1 m within and 6.1 to 7.6 m between rows and planting density ranges from 286 to 356 trees/ha (Crane, 1993). Interplanting with fruit crops such as papaya and banana is a common practice because these crops

17 provide an early income for growers and wind protection for young carambola trees (Crane, 1989). Shading. Carambola trees can survive under a wide range of light intensities (Marler et al., 1994). Shading is favorable for tree growth, and therefore, carambola trees can be successfully grown under shade cloth (allowing approximately 75% of sunlight through) used as a windbreak (Crane, 1994; Watson et al., 1988). Marler et al. (1994) exposed ‘Arkin’ seedling tree to approximately 25, 50, or 100% sunlight for 39 weeks and found that shading increased rachis length and leaf area but decreased leaflet thickness. Shading also reduced dark respiration, light compensation, and saturation points, but increased chlorophyll concentration and nitrogen-use efficiency. Under 100% sunlight, branches were more vertically orientated and stomatal density was higher than in shaded trees. Pruning. Pruning is an important cultural management practice used by carambola growers to improve light penetration, shape the canopy, limit tree size, and remove dead branches (Campbell, 1989; Crane, 1994; Green, 1987). Optimum tree height is about 3.5 to 4.5 m which facilitates manual harvesting (Campbell, 1989; Crane et al., 1991), improves equipment movement between rows, improves efficacy and penetration of foliar sprays, helps retain a bearing canopy throughout the tree, and reduces the potential for wind damage to the canopy (Crane et al., 1991). Pruning may stimulate flowering year-round (Núñez-Elisea and Crane, 1998, 2000), although Crane et al. (1991) found no consistent trend in flowering between pruned and non-pruned trees. Núñez-Elisea and Crane (2000) reported that early-season fruit production was increased by selective pruning and crop removal. They found that

18 pruning of 3- to 4-year-old branches to their main axes in early March resulted in flowering by mid-April and a crop by the end of June compared to non-pruned trees that produced mature fruit 4 to 5 weeks later. Pruning also reduced fruit wind-scar damage by 100%, whereas only 20% of non-pruned branches produced fruit free from wind damage. Núñez-Elisea and Crane (2000) reported that fruit removal from ‘Arkin’ carambola trees in November resulted in an average yield of 48 kg per tree on July, compared to an average yield of 5 kg per non-defruited trees. Irrigation. It is necessary to supply sufficient amounts of water to the soil to maintain optimum growth and yield of carambola especially during dry periods when rainfall does not meet tree water requirements. Drought before and after flowering can result in poor flowering, early fruit abscission, reduced yield and small fruit (Nakasone and Paull, 1998). Scheduling irrigation can be beneficial for commercial carambola production. However, little information is available about irrigation scheduling practices in carambola producing countries. General recommendations for irrigation scheduling were proposed in some regions of the world. During dry periods in northern Queensland and the Northern Territory of Australia, irrigation of 30 to 75 mm of water/tree per week was recommended for mature trees (Galán Saúco et al., 1993; Lim, 1996). In Malaysia where annual rainfall exceeds 2000 mm, Bookeri (1996) reported an increase in carambola yield (c.v. B17) with increasing irrigation rates of 4, 12, or 30 L/plant per day, but no water amount or irrigation frequency was recommended. In southern Florida, Crane (1994) recommended applying 33 mm of water per ha twice a week during dry periods

19 throughout the year. However, the optimum water application rates and frequencies have not been established. Microsprinklers are widely used to irrigate and fertigate carambola trees. Highvolume sprinklers are also used for irrigation and freeze protection (Campbell, 1989; Crane, 1993, 1994). Carambola irrigation in Florida is generally based on visual observations, since there are no established irrigation scheduling programs or water requirement data specifically for carambola trees in this area. Fertilization. A fertilizer application program for carambola trees should begin with soil analysis prior to planting (Galán Saúco et al., 1993). Soil and leaf sampling can be used to determine nutrient requirements of carambola trees (Galán Saúco et al., 1993). Young bearing carambola trees should receive 0.4 to 0.8 kg/ha per tree per year of NPK (11:12:17 to 15:15:15), while older trees (over 8 years old) may require 6 to 25 kg per tree per year (Nakasone and Paull, 1998). Fertilizer should be applied approximately every 3 months when trees are bearing fruit (Campbell, 1989). The Krome very gravelly loam soil in which carambola trees are grown in South Florida has low fertility and is often deficient in soluble iron, zinc, magnesium, and manganese due to the high pH (Campbell 1985; Campbell et al., 1985; Crane, 1993). Therefore, for adequate growth and yield, applications of chelated-iron, zinc, magnesium, and manganese are necessary to prevent deficiencies of these elements (Campbell, 1985, 1989; Crane, 1994; Knight, 1982). Fertilizer mixtures with 6-8% nitrogen, 4% phosphorous, 6-8% potash, and 3-4% magnesium are satisfactory for carambola trees (Crane, 1994). Young trees require light applications of NPK fertilizer (20-20-20) every 60-90 d until they are established. Two fertilizer applications per year are needed in

20 deeper soils and three applications per year in shallow soils. For young trees, fertilizer application rates are 400-500 g every 4-6 weeks during the rainy season or every 8 weeks during the dry season (Campbell, 1989). Growers in South Florida apply 500-600 kg/ha mixed fertilizers containing nitrogen, phosphorus, potassium and magnesium divided into 4-6 applications per year to mature orchards (Campbell, 1989). Chelated iron is applied annually at 50-100 g/tree, increasing to 300-400 g/tree as trees grow. Zinc and manganese are applied as inorganic salts, chelated compounds, or foliar sprays at a rate of 1.3-1.8 kg·l-1 of water (Campbell, 1989). Cultivars. Programs for selecting improved carambola cultivars have been established in Malaysia, Taiwan, Thailand, Florida and Hawaii (Nakasone and Paull, 1998). The Malaysian Agricultural Research and Development Institute (MARDI) was one of the most active centers for carambola selection and breeding. Some of the desirable characteristics for selections include: early season fruit maturity, abundant and regular production, ease of tree training, plasticity of growth, desirable fruit quality, and tolerance or resistance to biotic and abiotic stresses (Galàn Saúco et al., 1993). Several carambola cultivars have been selected and cultivated in different parts of the world (Table 2-1) (Galàn Saúco et al., 1993; Nakasone and Paull, 1998).

21

Table 2-1. Major carambola cultivars of economic importance in several major carambola production regions (Galàn Saúco et al., 1993; Nakasone and Paull, 1998). Major Cultivars Arkin, Golden Star, B10 Sri Kembangsaan, Kary Kaput, Ting Go, Demak Lang Bak, Juron Hong Hug, Far Dee Cheng Tsey B-2, B-10, B-17 Arkin, B-1, B-6, B-10, Jungle Gold Fwang Tung, Thai Knight Growing regions Florida, US Hawaii, US Indonesia Singapore China Taiwan Malaysia Australia Thailand

22 Several carambola cultivars have been selected for their adaptation to the South Florida climate. In addition, cultivars were screened for high fruit yields, good fruit quality, medium fruit size, yellow fruit color, ability of fruit to withstand damage during handling, length of fruit storage time, and marketing potential (Wagner et al., 1975). A study conducted by Crane (1989) showed that 89% of the carambola trees surveyed in southern Florida were 4-years-old or younger; 97% of these were cultivars with sweet fruit and 3% were cultivars with tart fruit. The dominant sweet cultivar has been ‘Arkin’ and the main tart type has been ‘Golden Star’ (Campbell, 1989; Crane, 1989; Knight and Crane, 2002). Other cultivars with very few commercial plantings in South Florida include ‘Kary’, from Hawaii, ‘B-10’, from Malaysia, ‘Fwang Tung’, from Thailand, and ‘Maha’ from Malaysia (Campbell, 1989; Crane et al., 1998; Knight and Crane, 2002). ‘Arkin’ is the most commonly planted commercial cultivar in South Florida to date because of its resistance to picking, packing and shipping damage, as well as its good marketability (Knight and Crane, 2002). The fruit are medium-sized, weigh between 90 and 200 g and are golden yellow at the early maturation stage, turning to yellow-orange when ripe. The ribs are thick and compact with a relatively large angle, and the edges of the wings are slightly rounded. The fruit has crisp texture and is juicy with a sweet flavor and relatively low acidity. Total soluble solid (TSS) content from July-harvested fruit was 4 to 8% (mean 6.8%) and that from November-harvested fruit was found to be 6 to 8% (mean 7.1%) (Crane et al., 1998). The fruit are also relatively insensitive to chilling injury during storage (Campbell 1989; Galán Saúco et al., 1993). Potential annual yield of carambola ranges between 4.5 to18 kg/tree when trees are 2 to 3 years old; 45 to 68

23 kg/tree at 5 to 6 years old; and 112 to 160 kg/tree at maturity between 7 to 12 years old (Campbell 1989; Crane 1994; Núñez-Elisea and Crane, 1998). Campbell (1985) selected and named ‘Golden Star’ in Florida in 1985. This cultivar has tart fruit that can attain a sweet flavor if allowed to ripen on the tree (Crane, 1994). Open pollinated seedlings of Golden Star are used primarily as rootstocks in Florida. Under south Florida conditions, Golden Star rootstocks are preferred to Arkin seedling rootstocks because they are well adapted to calcareous soils (Crane, 1994; Knight, 1982), moderately tolerant of flooded conditions (Joyner and Schaffer, 1989), and provide good tree anchorage (Crane, 1994). Environmental Stress Physiology of Carambola Information on the effects of environmental factors including temperature, drought, and flooding on tropical fruit crops such as carambola is less available in comparison to that for temperate fruit species. This is due in part on limited resources available for research and development in developing countries where tropical species are largely cultivated (Schaffer and Andersen, 1994). Among the adverse environmental conditions that affect carambola growth and yield are temperature extremes, flooding, wind, and water stress. Temperature Carambola trees are best adapted to hot lowland tropical climates but grow well in warm subtropical areas if not subjected to severe freezes (Campbell et al., 1985; Crane, 1994). Ideal temperatures for optimum growth and production of carambola are between 21 and 32oC (Nakasone and Paull, 1998; Ngah et al., 1989). High temperatures reduced yields by reducing fruit set due to lack of pollination. Temperatures of more than 30 to 35oC during flowering reduced pollen germination and

24 pollen tube growth (Salakpetch, 1987). Carambola leaflet petiolules are equipped with pulvini that allow leaflet movement. Under direct full sunlight, vertical leaflet movement helps reduce leaf temperature and increase photochemical efficiency of carambola leaves (Marler and Lawton, 1995). Marler and Lawton (1995) reported that maintaining a leaflet in a horizontal position for 3.5 h during midday under full sunlight reduced chlorophyll fluorescence and increased leaf temperature. Leaflet seismonastic movement also occurs in response to shaking of the tree canopy (Marler and Zozor, 1992). Low temperatures decrease carambola growth and freezing temperatures can lead to death of branches or trees (George and Nissen, 1994). Young trees can be killed if exposed to temperatures of -2.8 to -1.7 oC, while temperatures of -6.7 to -4.4 oC may kill large branches and mature trees (Campbell and Malo, 1981; Campbell et al., 1985; Watson et al., 1988; Crane, 1994). In Florida, carambola trees can be grown in warm locations along the southeastern and southwestern coasts (Campbell et al., 1985; Crane, 1994). The temperatures in south Florida are suitable for growing carambola about 9 to 10 months of the year, during which carambola trees grow vigorously and may grow vegetatively and reproductively at the same time. Carambola leaf production in Florida declines after mid-October with a heavy fruit load until December causing a sharp decline in tree carbohydrate reserves during the winter (Núñez-Elisea and Crane, 1998). The trees remain quiescent and partially defoliated until mid-March when refoliation occurs due to warm temperatures. George et al. (2000) reported that root mulching using organic and polyethylene mulches decreased defoliation during the winter months by increasing root temperature.

25 Wind Carambola is severely damaged by strong winds that can cause desiccation, dieback, defoliation and fruit scaring (Crane, 1993; Crane et al., 1994a; Campbell, 1985). Wind may reduce tree growth by reducing leaf gas exchange or by mechanical stress or damage (Marler and Zozor, 1992). Increasing wind speed from 0 to 2 m s-1 reduced net CO2 assimilation (A) of container-grown carambola trees by 22% and leaf area and dry weight production by about 50%. Wind-stressed carambola seedlings had less plant height, leaf area, dry weight, trunk cross-sectional area, a lower relative growth rate and leaf-area ratio, and lower A, and gs values than plants that were not wind-stressed (Marler and Zozor, 1992). In South Florida, carambola trees are frequently subjected to wind stress caused by strong winds, storms and hurricanes. Planting carambola trees in wind-protected orchards with man-made or natural windbreaks is a common practice in Florida. Commercially, carambola trees in Florida are grown within windbreaks that are either composed of trees with a rapid growth rate or fabric screens supported by posts and wires (Crane, 1993). After Hurricane Andrew in 1992, 93% of the carambola trees survived. Of the surviving trees, 13% were toppled, 4% were stumped (reduced to trunk), whereas 76% of the trees remained intact (Crane et al., 1993). Tree decline continued for 14 to 15 months after the hurricane. This decline was due to detachment of major roots and bark at the soil line (Crane et al., 1994a). Crane et al. (1994b) proposed several pre-hurricane and posthurricane practices to reduce the damage from hurricanes. Pre-hurricane practices include the use of grafted material and preparation of planting sites to increase tree rooting depth. After the hurricane, clearing the debris, resetting toppled trees, protecting trees from sunburn, and irrigating and fertilizing trees frequently hastened tree recovery.

26 Flooding Carambola is considered a moderately flood-tolerant tree (Crane, 1994; Joyner and Schaffer, 1989). However, continuous flooding for up to 18 weeks resulted in reduced A, transpiration (E), and gs (Joyner and Schaffer, 1989). Productivity of mature trees was reduced under flooded conditions (Campbell, 1989) because of reduced leaf, shoot, and root dry weights (Joyner and Schaffer, 1989). However, after flooded trees were unflooded, trees were able to recover from continuous and intermittent flooding and A, E, and gs increased to pre-stress levels (Joyner and Schaffer, 1989; Ismail and Noor, 1996b). Drought and water stress Carambola trees grow best in tropical lowlands with uniform rainfall distribution throughout the year (Campbell and Malo, 1981; Samson, 1980). However, trees can also grow well in humid, subtropical areas (Campbell and Malo, 1981). In some dry areas, carambola trees are very tolerant of seasonal drought periods once they are well established (Galán Saúco et al., 1993). The optimum rainfall for maximum production in Malaysia is 2500 mm annually (Ngah et al., 1989), while rainfall between 1500 to 3000 mm is suitable for production in Australia (Galán Saúco et al., 1993). Locations with annual rainfall above 1800 mm in Australia produced fruit with better flavor and quality compared to trees in areas with less rainfall (Morton, 1987). Good quality fruit have been produced in the Canary Islands with annual rainfall and irrigation of 800 mm (Nakasone and Paull, 1998). Average annual rainfall in south Florida where carambola trees are grown is 1500 mm, two-third of which occurs from May to November. Vegetative growth. Shoot expansion in plants is greatly affected by water stress and therefore can be an indicator of water stress (Hsiao, 1973; Hsiao and Bradford, 1983). Vegetative growth of carambola decreases with increased water stress (Ismail and

27 Awang 1992; Ismail et al., 1994, 1996), particularly when root growth is restricted (Ismail and Noor, 1996a). An adaptive mechanism to water stress is greater allocation of dry matter to the roots which results in a high root to shoot ratio in water-stressed plants (Ismail and Awang 1992; Ismail et al., 1996). Stressed plants, however, continue to produce new leaves, although at a slower rate than nonstressed control trees (Ismail et al., 1996). Leaves wilted 10 d after withholding water from 2-year-old ‘Fwang Tung’ and ‘Thai Knight’ carambola trees (Salakpetch et al., 1990). Leaf wilting coincided with midday leaf water potentials (ΨL) of approximately -2.0 MPa. Five days later, some leaves became yellow and abscised. Upon rewatering, leaves appeared normal (nonwilted) and ΨL returned to previous levels (-0.6 MPa) 2 to 3 d later. Trunk diameter or tree girth can be used to evaluate the effect of long-term and short-term stress on fruit trees. Bookeri (1996) found no significant difference in girth between irrigated and non-irrigated (rainfed) young carambola trees because sufficient rainfall occurred following periods of drought. Reproductive growth and yield. The response of carambola tree flowering to water stress depends on the time of stress. Flowering of carambola trees is more strongly influenced by water stress and cultivar than by diurnal temperature variation and photoperiod (Salakpetch et al., 1990). Ismail et al. (1996) reported that water stress induced early flowering in container-grown young carambola trees by approximately 6 weeks. Production of new flowers continued in water-stressed trees and there were more flowers in water-stressed than in well-watered trees. However, flowering was reduced when water stress occurred during flowering or fruit set. In addition, prolonged severe water stress may reduce flowering of carambola trees. In a heated glasshouse experiment,

28 Salakpetch et al. (1990) found that cyclical or continuous (-2.0 MPa midday ΨL) water stress for 3 to 4 weeks inhibited flowering of ‘Thai Knight’ carambola. Flowering of carambola trees is also influenced by root growth. Flower initiation was hastened by restricting root growth even under well-watered conditions (Ismail and Noor, 1996a). Although tree growth was not affected by drought, yield was significantly increased in irrigated compared to rainfed trees (Bookeri, 1996). The increase in yield was attributed to a 17% increase in fruit size, although fruit number was not affected by irrigation (Bookeri, 1996). Physiological processes. Leaf water potential decreases when water lost via E is greater than water uptake (Salisbury and Ross, 1985). Leaf water potential of carambola trees declined with increasing water stress (Ismail and Noor, 1996a; Ismail et al., 1994, 1996). Ismail et al. (1994) carried out an experiment on 6-month-old, container-grown ‘B17’ carambola trees grafted onto B2 rootstock in a greenhouse and observed that the mean midday ΨL of water stressed trees was -2.20 to -0.35 MPa lower than that of nonstressed trees. Ismail and Noor (1996b) found that ΨL (measured 4 h after sunrise) of flooded carambola trees declined progressively to -1.3 MPa 21 d after trees were flooded compared to -0.80 MPa for non-flooded trees. Ismail et al. (1994) studied leaf gas exchange of 6-month-old, container-grown carambola trees grown in a greenhouse at mean ambient temperatures of 29.4 ± 5.6 oC subjected to water stress. The mean A of non-stressed trees was 3.13 µmol.m-2.s-1 compared to 0.24 to 1.83 µmol.m-2.s-1 for the water stressed trees. Stomatal conductance was two to three times lower in water-stressed trees than in control trees. In another study of carambola trees in a glasshouse, Ismail and Awang (1992) observed that A was only

29 reduced when ΨL fell below -0.85 MPa and that plants were photosynthesizing 14 d after withholding water. They also found that gs, respiration, and chlorophyll content were only reduced 7 d after withholding water. Reduction in A could be attributed to reduced gs. Ismail et al. (1994) reported that A was reduced when stomatal resistance exceeded 3.5 s cm-1. They concluded that a photosynthetic photon flux (PPF) higher than 1000 µmol.m-2.s-1 reduced A of stressed and non-stressed plants due to photoinhibition. Care must be taken when measuring leaf gas exchange of carambola trees because leaves are sensitive to mechanical stress caused by the leaf cuvette clamping onto the leaf during gas exchange measurements. This mechanical damage reduced gs of carambola leaves (Marler and Mickelbart, 1992). Water Stress Lack of water is one of the most limiting environmental factors for crop production in the world. Plants require large amount of water to produce their biomass, ranging from several hundred to 2000 g of water per g of dry matter produced (Hsiao, 1993). Thus, plant water status is perhaps the most important factor that must be controlled to obtain high yields of good quality horticultural crops (Jones, 1990). Plant responses to water stress depend on the quantity of water lost, the rate of loss, and the duration of stress (Bray, 1997). Water stress can reduce vegetative growth, flowering and fruit growth. Hanan (1972) defined water stress as the difference between water supply and demand. A water deficit occurs when a plant’s E exceeds its water uptake and is a component of other stresses including drought, salinity, and low temperature (Bray, 1997). Water stress is therefore caused by any restriction in supply or resistance to flow through the plant when demand for water increases. Halevy (1972) stated that water stress is a result of an imbalance between three components that determine plant water status: water absorption,

30 translocation and loss. Lakso (1985) argued that the relative sparsity of roots in fruit crops leads to low root hydraulic conductivity and thus under high evaporative demand, low plant Ψ develop. Therefore, in fruit trees, daily plant Ψ is controlled more by the evaporative demand than by soil water tension. Thus, for fruit trees, plant water status may be a better determinant than soil water content for scheduling irrigation to obtain the desired growth and yield. Water deficits impact many physiological and developmental processes that affect fruit production, including growth (cell division and cell expansion), and gas exchange (stomatal aperture, and photosynthetic and respiratory enzymes) (Jones et al., 1985). The response of fruit trees to water stress varies between species and varieties within the same species. Germanà and Sardo (1996) reported variable responses to water deficit for 41 citrus cultivars grown under nearly identical vapor pressure deficits (VPD) and PPF. They cited morphological and functional differences among the leaves as factors influencing water use efficiency (WUE) where larger leaves with higher stomatal densities, had higher A with no significant difference in E compared to trees with smaller leaves. Mild water stress increased water use efficiency of apple (Davies and Lakso, 1979; Flore et al., 1985), cherry (Flore et al., 1985) and peach (Chalmers et al., 1984; Girona et al., 1993). Turner (1979; 1986) separated the mechanisms of plant adaptation to water deficits into three categories: 1) Drought escape: the ability of a plant to complete its life cycle before serious soil and plant water deficits develop. This can be achieved by rapid phenological development and developmental plasticity.

31 2) Drought tolerance with low plant water potential: the ability of a plant to endure periods without significant rainfall and endure low tissue water status, i.e. dehydration tolerance (Kramer, 1980). The mechanisms involved in this process include maintenance of turgor by osmotic adjustment, increase in cell elasticity, a decrease in cell size, and desiccation tolerance through protoplasmic tolerance. 3) Drought tolerance with high plant water potential: the ability of a plant to endure periods without significant rainfall while maintaining a high plant water status, i.e. dehydration postponement (Kramer, 1980). This can be achieved by reduction of water loss through an increase in stomatal and cuticular resistance, a reduction in radiation absorbed, and/or a reduction in leaf area, and maintenance of water uptake through increased root density and depth and increased hydraulic conductance. The root system of fruit crops affects the rate of water stress development. A slow rate of onset of stress may allow for development of adaptive mechanisms such as osmotic adjustment, decreasing leaf area, abscission of leaves, leaf folding, rolling or reorientation of leaves, and an increased root growth rate (Flore and Lakso, 1989). Lankes (1985) found that drought tolerance of apple leaves and fruit varied depending on the time of season or stage of development. In the early stage of development, leaf A and E responded in a pattern of ‘drought tolerance at high water potential’ and later in the season, the response pattern was ‘drought tolerance at low water potential’. Osmotic adjustment is an example of avoidance at the cellular level as a low rate of water loss may allow acclimation to water deficit (Bray, 1997).

32 Effect of Water Stress on Vegetative Growth Root growth and distribution Plants often respond to water stress by a shift in new assimilate distribution to support the growth of roots at the expense of the shoot (Mansfield and Atkinson, 1990). Enhanced root growth under water stress is due to osmotic adjustment in root cells (Hsiao, 1973). Shallow roots of avocado can grow deeper than 1.2 to 1.5 m in welldrained sandy soils or under restricted water supply compared with other fruit trees (Lahav and Whiley, 2002). The root to shoot ratio was 3.5 times higher in water-stressed kiwifruit vines compared to well-watered vines (Chartzoulakis et al., 1993). In carambola trees, the root to shoot ratio was proportional to the duration of water stress (Ismail et al., 1994). Leaf area Leaf growth is known to be very sensitive to water stress. Leaf area is often reduced when trees are subjected to water stress. In deciduous fruit trees, when the canopy is incomplete at the beginning of the growing season and intercepts only part of the incident radiation, even mild water stress can slow leaf growth (Hsiao, 1993). A reduction of 30% (70% of total ET) of irrigation water reduced leaf area in walnut (Cohen et al., 1997). A reduction in leaf area reduced A leading to reduced biomass accumulation. Chartzoulakis et al. (1993) reported that decreased growth of kiwifruit induced by water stress was a result of a reduction in A and preferential partitioning of photosynthate to the roots and thus affected leaf area development. Elongation of leaves of container-grown, 6-month-old carambola trees was reduced as a result of water stress (Ismail et al., 1994). Leaf area of trees was reduced by 60% when water was withheld for

33 1 week and 90% when water withheld for either 2 or 4 weeks before restoring soil to field capacity (Ismail et al., 1994). Trunk growth Responses of stem diameter to changes in plant water status and the development of instruments that automatically measure these changes have lead to the use of trunk diameter as a method for irrigation scheduling for several fruit crops (Hilgeman, 1962; Higgs and Jones, 1984; Huguet et al., 1992; Simonneau et al., 1993). A decrease or cessation of daily stem diameter growth was the first indication of water stress in fieldgrown apple and peach trees. If trees received a sufficient amount of water, maximum daily stem shrinkage was a versatile indicator of E (Huguet et al., 1992). The change in diameter of a living stem over 24 h can be described by measurements of maximum daily shrinkage (MDS) which is the maximum diameter usually observed in the early morning and the minimum diameter usually observed in mid-afternoon, and daily evolution (DE) which is the total change in stem diameter over a 24-h period beginning at dawn (Huguet et al., 1992; Katerji et al., 1994). Maximum daily stem shrinkage is affected by environmental factors affecting plant E, e.g. soil water content and potential evapotranspiration. Trunk diameter fluctuations can be measured with linear variable differential transformers (LVDT) (Huguet et al., 1992). Trunk diameter variations were a good indicator of water stress in walnut (Cohen et al., 1997), peach (Garnier and Berger, 1985; Huguet et al., 1992; Girona et al., 1993), avocado (Lahav and Kalmar, 1977), cherry (Kozlowski, 1968), citrus (Hilgeman, 1962), and apple (Huguet et al., 1992). Huguet et al. (1992) observed two types of specific daily shrinkage patterns: (1) a ‘peach-tree’ pattern, where maximum daily shrinkage increased as the severity of water stress

34 increased; and (2) an ‘apple-tree’ pattern, where maximum daily shrinkage was smaller under severe water stress than that of well-watered trees. Shoot growth Reductions in extension shoot growth and leaf expansion are the first visible responses of crop plants to water stress (Hsiao and Acevedo, 1974; Lakso, 1985). In grapevines, shoot growth was found to be more sensitive to water stress than fruit yield (Nagarajah, 1989). In crop plants, defoliation and restricted leaf growth are water stress adaptations aimed at reducing E and conserving water (Nagarajah, 1989). A decrease in shoot growth caused by a water deficit is a result of a reduction in both A and photosynthate partitioning to the shoot in favor of root growth (Chartzoulakis et al., 1993). Carambola (Ismail et al., 1994) and avocado (Lahav and Whiley, 2002) shoot growth was significantly reduced under water stress conditions. Effect of Water Stress on Reproductive Growth and Yield Flowering and fruiting Water status of flowers and their response to water stress depends on their stage of development. Halevy (1972) stated that young flowers of gladioli, up to about 2 d prior to anthesis of the first floret, were more sensitive to drought than old leaves because they were the first to wilt. Water relocation was reversed just before anthesis when water was translocated to the flowers from other organs. Hsiao (1993) suggested that during severe water stress at the time of anthesis, pollination can be prevented and the number of reproductive sinks for assimilates is reduced. However, mild to moderate water stress before and during anthesis can enhance the partitioning of assimilates toward reproductive sinks. Increased partitioning of assimilates to the reproductive sinks then promotes embryo development which in some species can result in early maturation of a

35 portion of the fruit. Menzel et al. (1995) found that before anthesis in lychee trees, tree water status was not related to extension growth of floral panicles or leafy shoots. In contrast, they found that no vegetative shoots were initiated after fruit set in droughtstressed trees when ΨL declined to –2.5 MPa. A reduction in the number of fruit as a result of moderate to severe water stress is often attributed to an inhibition of pollination. However, more likely causes of fewer fruit as a result of water stress are a reduced number of fruiting sites associated with the smaller plant size and abortion of younger fruit (Hsiao, 1993). Water deficits reduced initial fruit set of lychee by 30% and final yield by 70% as a result of fruit splitting (Menzel et al., 1995). ‘Valencia’ orange had delayed bloom and lower fruit set when irrigated at soil water tension of 70 kPa compared to trees irrigated at 45 kPa or below 20 kPa (Davies and Bower, 1994). Predawn leaf water potentials of –1.7 to –3.5 MPa resulted in reduced growth but did not induce flowering in avocado and lychee compared to trees with leaf water potentials of –0.4 to –0.7 MPa (Chaikiattiyos et al., 1994). Similar responses were observed for mango when predawn relative water content was 90-93% compared with 97% or above in control trees. Schaffer et al. (1994) stated that a dry period preceding flowering is essential for reliable mango production in the tropics. In lemon trees, flowering is mainly determined by water stress over a range of daytime temperatures from 18oC to 30oC. More flowers were produced by water-stressed lemon trees in comparison to well-watered trees (Barbera and Cascio, 1985; Chaikiattiyos et al., 1994). Similar observations were made for ‘Tahiti’ lime by Southwick and Davenport (1986, 1987).

36 Yield Yield losses in water stressed plants can also be attributed to a reduction in inflorescence initiation, fruit set, and increased fruit abscission (Nagarajah, 1989). The effect of water stress on yield varies among tree species and cultivars. Reduction in yields due to water stress has been extensively studied in several fruit crops in temperate, tropical and subtropical climates. Water stress reduced fruit size and yield of temperate, tropical, and subtropical fruit trees including apples (Failla et al., 1992), tangerines (Huang et al., 2000), walnut (Cohen et al., 1997), grapes (Nagarajah, 1989), apricot (Torrecillas et al., 2000), lychee (Batten et al., 1994; Menzel et al., 1995), and avocado (Lahav and Whiley, 2002). In an irrigation experiment in Miami-Dade County, Florida, fruit of ‘Tahiti’ lime matured earlier (before 1 June) when one third (high) or two thirds (moderate) of available soil moisture had been depleted compared to trees irrigated 2 d following ‘midday wilt’ with 22.86 mm of irrigation water (Krome et al., 1970). However, total annual yield was not significantly different among lime trees subjected to these three irrigation treatments. Fruit quality Water stress affects several fruit quality factors. Nagarajah (1989) reported that mild water stress during the ripening stage of grapevines increased the concentration of sugars and anthocyanins and decreased acid and pH levels. Moderate water stress improved fruit quality of prunes (Shackel et al., 2000). Total soluble solids (TSS) of satsuma mandarin trees increased with deficit irrigation of -30, -50, -70, or -100 kPa as determined with tensiometers (Peng and Rabe, 1998). Frequent microsprinkler irrigation at low soil tension (10-15 kPa) also resulted in increased fruit size and increased TSS of Florida grapefruit when compared to high soil tension (34-45 kPa) with less frequent

37 irrigation (Boman, 1996). Domingo et al. (1996) subjected ‘Fino’ lemon trees to three irrigation levels: 100% ETc all year, 25% ETc all year but 100% ETc during fruit growth, and 100% ETc all year but 70% ETc during fruit growth. They found no significant differences among treatments in TSS, acidity, ascorbic acid glucose, fructose, and sucrose levels. Avocado oil content was 10% higher in trees irrigated at 7-d intervals with 889 mm per year than in trees irrigated every 28 d with 594 mm of water per year (Lahav and Kalmar, 1977). Effect of Water Stress on Physiological Processes Net CO2 assimilation and transpiration Net CO2 assimilation and E are processes that involve gas and water vapor exchange through the stomata. To study the effect of water stress on these processes, it is recommended that they be examined together (Lakso, 1985). The relationship between A and gs is different than that between E and gs (Lakso, 1985; Schulze, 1986). Under a constant vapor pressure gradient, E is linearly correlated with gs, whereas A is hyperbolically correlated. Under low water stress, gs may be very high and partial stomatal closure may reduce E with little effect on A (Lakso, 1985). Both A and E are reduced equally, however, under low humidity or progressive water stress that induces stomatal closure (Lakso, 1985). The difference in vapor pressure between leaf and air, i.e. vapor pressure gradient, is very important at both the leaf and canopy levels, because it drives E and directly affects gs and A (Schulze, 1986). Water uptake plays an important role in plant A. There is less resistance to water efflux than CO2 influx into leaves through the stomata (Hsiao, 1973, 1993). Therefore, large amounts of water are lost through E as a consequence of stomates opening to allow entry of carbon dioxide into the leaves. Transpiration varies in the morning from day to

38 day depending on variability in evaporative demand. Thus, early afternoon measurements, when E is low but stable, are usually recommended (Braun et al., 1989). During the development of water stress, stomatal closure is the dominant mechanism restricting water loss in mesophytes (Hsiao, 1973). This affects A directly by restricting the CO2 supply. The decrease in gs is therefore an indicator of plant water stress (Braun, et al., 1989; Nagarajah, 1989). However, it should only be measured on clear days, to exclude effects of rapidly changing light and heat conditions (Braun et al., 1989). Severe water stress can affect the photochemical and enzymatic reactions of A (Nagarajah,1989). Reduction in A in response to water stress has been reported in many fruit crops such as satsuma mandarin (Peng and Rabe, 1998), kiwifruit (Chartzoulakis et al., 1993), lychee (Menzel et al., 1995), ‘Valencia’ orange (Davies and Bower, 1994), peach (Girona et al., 1993), pistachio (Palma et al., 1997), apple (Sritharan and Lenz, 1989), carambola (Ismail et al., 1994), lemon, and lychee (Chaikiattiyos et al., 1994). Stomatal conductance Plants exhibit two basic stomatal-related responses to water stress which help to alleviate the stress (Mansfield and Atkinson, 1990). The first response is a shift in distribution of new assimilates from shoots to roots to support root growth at the expense of shoot growth. This increases root area and the capacity of the root to obtain water and restricts leaf expansion to minimize water loss through E. The second response of plants to water stress is an adjustment in the degree of stomatal opening to inhibit water loss by E. A reduction of A and E of grapevines (Nagarajah, 1989), and kiwifruit (Chartzoulakis et al., 1993) among other crops was attributed to stomatal closure caused by reduced ΨL. Flore et al. (1985) found that stomata of well-watered apple, cherry, and blueberry plants were more responsive to vapor pressure deficit and less responsive to the CO2 gradient

39 than stressed plants. In peach, leaf gs was reduced in water-stressed trees compared to well-watered trees (Tan and Layne, 1991). Stomata of most plants close in response to an increase in the VPD between the leaf and the surrounding air, which influences the turgor of stomatal guard cells (Lakso 1985; Mansfield and Atkinson, 1990). Other factors that play a role in stomatal closure under water stress conditions include carbon dioxide (CO2) and leaf abscisic acid (ABA) concentration (Flore et al., 1985). The level of ABA in the leaf increases during water stress in many fruit crops including apples (Davies and Lakso, 1978) and grapes (Nagarajah, 1989). Abscisic acid is a major signaling phytohormone for water stress that is synthesized in leaves and in roots (Bray, 1997; Jackson, 1997; Mansfield and Atkinson, 1990; Schulze, 1986; Seeley, 1990). Abscisic acid concentration increased linearly with decreased leaf turgor in apple seedlings (Davies and Lakso, 1978). Turgor-controlled ABA synthesis is essential for regulating water loss under water stress. At the early stage of water stress, ABA is translocated in the E stream through xylem vessels from the roots to the leaves to bring about stomatal closure, and more ABA is synthesized as water stress progresses (Mansfield and Atkinson, 1990; Comstock, 2002). Stomatal closure in response to water stress is vital in perennial fruit crops because plant Ψ is dependent on E (Lakso, 1985). Leaf and stem water potential Water potential is a measure of the chemical potential of water and is essential in determining the direction of water flow (from higher to a lower potential), as in the E stream (Jones et al., 1985; Jones, 1990; Plaut and Meiri, 1994). Water potential has been used as a measure of plant water status and consists of osmotic potential (Ψo) pressure or turgor potential (Ψp), matric potential (Ψm), and gravitational potential (Ψg) (Jones et al.,

40 1985). Gravitational potential (Ψg) increases by 0.01 MPa per 1 m of plant height, therefore it is less than 0.05 MPa for most trees and is usually ignored when assessing a tree’s water status (Jones et al., 1985). A reduction in osmotic potential (Ψo) due to increased solute content or decreased water content (dehydration) is a result of water stress (Plaut and Meiri, 1994). Turgor potential (Ψp) is considered a better measure of plant water status than osmotic potential (Ψo) (Hsiao and Bradford, 1983) and its reduction as a result of water stress can be detrimental to physiological processes related to growth. However, direct measurement of plant turgor pressure is difficult under field conditions (Zimmermann, 1978). Leaf xylem water potential as an indicator of plant water status can be determined using a pressure chamber (Scholander et al., 1965). Different physiological processes vary in their sensitivity to reduced ΨL caused by water stress. Cell expansion can be inhibited at a ΨL of approximately –0.4 to –0.8 MPa (Plaut and Meiri, 1994), while photosynthetic CO2 fixation is much less sensitive to changes in ΨL and can be reduced at ΨL of –1 to –2 MPa (Hsiao and Acevedo, 1974). In citrus, midday Ψ is stabilized by effective stomatal control of leaf water status under a wide range of evaporative demands (Levy, 1980). For plants that have been moderately water stressed, midday ΨL can be higher than that of well-watered plants due to stomatal closure (Levy, 1980). Similar effective stomatal control of ΨL has been reported in apple (Jones et al., 1983). In grapevines, Nagarajah (1989) indicated that water stress reduced both water and turgor potential. The reduction in ΨL induced stomatal closure and increased ABA levels in the leaves. Stomatal closure reduced A and E (Nagarajah, 1989). Water potential of lychee trees declined to –2.6 to –2.8 MPa under water stress where soil water content reached minimum of 7.6% compared to 15.5% for well watered soils

41 (Menzel et al., 1995). Predawn ΨL differences were more significant among water deficit treatments than midday ΨL in peach trees (Girona et al., 1993). In carambola, Ismail et al. (1994) reported a decrease in ΨL and relative water content with increased water stress. They also found that ΨL at predawn was – 0.65 MPa and – 0.9 MPa at midday, while ΨL after 4 weeks of withholding water was – 2.52 MPa at midday. With decreasing soil water content, water uptake in trees during the afternoon and night becomes increasingly less resulting in low pre-dawn ΨL (Braun et al., 1989). Since Ψp = Ψ – Ψo, during stress, as Ψ decreases, a positive Ψp is maintained by a reduction in Ψo (Nagarajah, 1989). This adaptive response by which Ψo is reduced in response to water stress is called osmotic adjustment (Begg and Turner, 1976; Hsiao and Bradford, 1983). Maintaining positive Ψp is essential for the continuation of many processes in the plant such as cell expansion, stomatal movement, and enzymatic reactions (Nagarajah, 1989). Osmotic adjustment may occur in different parts of the plant and varies depending on the level of stress, species and variety, season and growth stage. In grapevines for example, osmotic adjustment takes place in leaves or shoot apices but not in fruit or root apices (Nagarajah, 1989). In apple fruit, osmotic adjustment occurred during the enlargement phase but not in the cell division phase of fruit growth and development (Failla et al., 1992). As stress developed in apple, sorbitol, glucose, and fructose concentrations in the leaves increased, while sucrose and starch levels decreased significantly, strongly suggesting that sugar alcohol and monosaccharides are the most important osmotica for osmotic adjustment in apple (Wang and Stutte, 1992).

42 Irrigation Scheduling of Fruit Trees Optimizing water applications to fruit orchards may increase water conservation, reduce production costs, and increase plant growth and yield. Irrigation scheduling is especially important in horticultural crops because net returns are normally higher than those of other crops (Fereres, 1997). According to Hillel (1998), the main issue with irrigation management is to determine the frequency, quantity, and timing of irrigation to optimize crop growth and productivity. Fereres (1997) stated that irrigation scheduling of horticultural crops is essentially reduced to predicting the amount of water required to meet the crop demand. The basis for predicting a crop’s water requirements is an estimation of crop ET (Fereres, 1997). Measurement of soil water content and soil matric potential provide an index of the rate at which water is taken up by the plant or lost from the root zone. Soil water content or potential is therefore most useful in conjunction with other information about the soilplant-atmosphere system (Campbell and Campbell, 1982). Water extraction from the soil is dependant on plant properties that determine the plant Ψ at which a particular plant species can continue to grow and extract water from the soil (Jones, 1990). The majority of subtropical and tropical fruit crops in South Florida are irrigated at rates and frequencies based on experience and observations of crop growth and yield rather than on quantitative scientific information (Schaffer, 1998). Li (2000) suggested several methods of irrigation scheduling for tropical fruit orchards in South Florida. These included irrigation during flowering at soil tension (determined with tensiometers) of 10-15 kPa and at 15-20 kPa at other times. He also suggested irrigating midway between field capacity and the onset of stress as determined with an EnviroSCAN

43 multisensor capacitance probe system and associated software. Li (2000) also suggested indirect methods of irrigation scheduling based on the following equation: A = ET·L·W·15.8, where: A is the amount of water required (L/tree per day), ET is evapotranspiration (mm·d-1), L is the spacing between trees within the row (m), and W is the spacing between rows (m). Although the previously suggested methods provide a means for estimating irrigation amount and timing, they do not take into account the variability between fruit tree species and cultivars, growth stage, or the response of trees to soil moisture deficit. The water refill point, which is the lowest possible soil water content with no decrease in yield or fruit quality, varies among different tree varieties, rootstocks, soils and seasons (Braun et al., 1989). It is often beneficial to use both soil and plant factors for irrigation scheduling. An integrated approach utilizing soil and plant factors suggested by Buss (1989) included soil surveying to determine soil properties and available soil moisture content and crop factors that included crop type, canopy size, rooting width and depth, and crop density. Irrigation scheduling was then planned from a combination of available water data, crop water extraction rates, and the irrigation system layout. Irrigation scheduling based on plant water status The effect of several factors that control water status were summarized by Jones (1990) and divided into four groups: (a) the soil Ψ and the factors involved in its control. These include rainfall and water use, soil texture and depth, and root distribution and activity; (b) crop evapotranspiration (ET), historic and current; (c) hydraulic conductances within the plant that are important for controlling water; (d) the relationships between plant Ψ and other measures of plant water status (i.e. turgor

44 pressure, osmotic potential, or water content) which can be affected by processes such as osmotic adjustment, cell wall elasticity and mesophyll structure. Several techniques used to schedule irrigation including destructive and nondestructive, and discrete and continuous water status measurements in plant populations, individual plants, or plant organs. Methods of irrigation scheduling based on plant responses to soil water include the use of remote sensors such as radiation sensors and infrared sensors, and measurements of trunk diameter, ΨL, and stem water potential (ΨS), and sap flow. Irrigation scheduling based on atmospheric or soil variables include basing irrigation on ET or soil water status monitored with devices such as tensiometers, neutron probes, or capacitance probes. Several physiological variables are used as indicators of plant water status. Among the most frequently used is ΨL (Hsiao, 1990). Plant water potential. Plaut and Meiri (1994) defined Ψ as the quotient of the chemical potential of water and its partial molal value (18 cm3 mol-1). The chemical potential of water is a qualitative expression of the free energy of water and is influenced by three factors: concentration, pressure, and gravity, all expressed as the difference between the chemical potential at a given state and its value at a standard state (Plaut and Meiri, 1994). The Ψ concept satisfies two main functions (Plaut and Meiri, 1994): (a) it is the driving force of water movement from the soil to the root and it controls the direction of water flow across cell membranes; and (b) it is a measure of plant water status that can be used as a plant stress indicator and for irrigation scheduling. Naor et al. (1999) found that total yield and fruit size of nectarines grown in a semiarid zone were more highly correlated with midday ΨS than with soil Ψ. Similar results were reported for apple (Naor et al., 1995) and pear (Shackel et al., 1997). A low correlation between yield of nectarine

45 and soil Ψ was attributed to the typically high variability in tensiometer readings (Naor et al., 1999). Several techniques have been developed to determine plant Ψ. Among the most commonly used are the pressure chamber and thermocouple psychrometer (Turner, 1981; Phene et al., 1990). A pressure chamber is used to determine plant ΨL (Scholander et al., 1965). With this technique, the leaf or branch is cut and enclosed in a steel pressure chamber with the cut end protruding through a rubber stopper which is used to seal the chamber. The pressure in the chamber is gradually increased until the sap appears at the end of the xylem vessels. After the pressure is recorded, it is released through an outlet valve and the sample is removed (Turner, 1981; Phene et al., 1990; Shackel et al., 1997). It is often necessary to use a microscope or hand lens to observe the point at which sap first starts to exude from the xylem. The entire procedure can be completed in 2 min for each leaf, depending on the dehydration level of the tissue. This allows for the measurement of about 20 to 30 samples in an hour (Turner, 1981). To avoid water loss, leaves should be enclosed in a plastic bag prior to excision (Turner and Long, 1980). Stem water potential can also be measured with the pressure chamber by covering the leaves prior to measurement for a minimum of 30 min (Begg and Turner, 1970). Shackel et al. (1997) found that in prune trees, ΨS was more sensitive than ΨL as an indicator of progressive water stress and suggested its use as a guide for making irrigation decisions. Irrigation scheduling based on soil water status Irrigation scheduling based on grower observations of soil moisture is an ancient practice and is still the major method of irrigation timing used by fruit growers. Soil water is usually measured as water content (mass or volume fraction) or as soil Ψ (i.e., the energy required to remove water from the soil). Measurements of soil water content

46 can be direct (gravimetric or volumetric methods) and indirect that involve installation of sensors that measure soil water content, such as capacitance probes and time domain reflectometry, or portable equipment such as the neutron scattering probe, or measuring soil matric potential using tensiometers. Gravimetric or volumetric method. The gravimetric or volumetric method is a direct measurement of soil water content. The percent of water is measured per mass or volume of soil using the ratio of dry weight to wet weight of the soil sample (Phene et al., 1990). This is considered a destructive method since it involves soil sampling from the field (Campbell and Campbell, 1982). Volumetric or gravimetric methods require drying the soil, and therefore may take too long for making decisions about when to irrigate which often need to be made instantaneously. Soil matric potential (tensiometers). Tensiometers were developed in the 1920s (Richards, 1942) and measure soil suction or matric Ψ rather than soil water content (Muñoz-Carpena et al., 2002; Phene et al., 1990; Smajstrla and Harrison, 1998). Soil suction or matric potential is defined as the amount of water held by capillarity against the force of gravity and is often measured in units of pressure such as kPa (MuñozCarpena et al., 2002). Soil water tension is also a measure of the energy required for plants to extract water from the soil (Smajstrla and Harrison, 1998). A soil water retention curve must be established to determine the soil water content that corresponds to the water matric potential in order to estimate soil water content with a tensiometer. The main components of a tensiometer include a porous ceramic cup, a water column, and a pressure-measuring device (usually a vacuum gauge). Tensiometers are filled with water then calibrated with a manual pump or vacuum chamber (Smajstrla and Pitts, 1997)

47 to remove air from the water column, to check for leakage, and to ensure accurate gauge readings. There must be good contact between the ceramic cup and the surrounding soil (Phene et al., 1990) to insure the accuracy of the reading. Therefore, for accurate tensiometer reading in porous, rocky soil such as Krome very gravelly loam, it is essential to apply soil slurry around the ceramic cup when tensiometers are installed (Núñez-Elisea et al., 2001). Tensiometers are relatively simple to use and inexpensive (Campbell and Campbell, 1982). A disadvantage of using tensiometers for determining soil water content is that they are limited to measuring soil matric potential up to 40 kPa for lowtension tensiometers and 80 kPa for high-tension tensiometers (Campbell and Campbell, 1982; Phene et al., 1990). At higher tensions, air enters the tensiometer cup (Phene et al., 1990) and makes the readings inaccurate. Moreover, tensiometers require frequent servicing to function properly. Also, they measure soil matric potential within a very limited volume; therefore, several tensiometers are needed for a reliable measurement (Campbell and Campbell, 1982) of soil water matric potential. In Krome very gravelly loam soils in southern Florida, soil water tension readings from tensiometers were more highly correlated with volumetric water content when tensiometers were installed at a depth of 10 than at 30 cm (Núñez-Elisea et al., 2001). Lower correlation at 30 cm depth was due to the presence of more rock fragments larger than 2 mm at the 30 than at the 10 cm depth (Núñez-Elisea et al., 2001). However, air entry into tensiometers at tensions greater than 20 kPa was observed in Krome very gravelly loam soils (Al-Yahyai et al., 2003), where tensiometer readings did not reflect changes in soil water content (Muñoz-Carpena et al., 2002).

48 Neutron probe. Neutron probes measure soil volumetric water content. The main component of a neutron probe is a fast neutron source encased in a protective shield and an electronic counting scaler which are connected by an electric cable that is also used to lower the probe into an access tube to determine soil moisture in the soil profile (Chanasyk and Naeth, 1996). Neutrons with high energy are emitted by a radioactive source, such as americium 241/beryllium, into the soil and are slowed down by elastic collisions with nuclei of atoms, primarily hydrogen in the soil (Gardner and Kirkham, 1952). The density of the resultant cloud of slow neutrons is a function of the soil moisture content (Chanasyk and Naeth, 1996). The number of fast neutrons that are slowed is measured as a ‘count rate’ per unit time (Gardner et al., 1991). The count rate is converted to volumetric water content using a calibration curve. A polyvinyl chloride (PVC) or aluminum access tube is inserted vertically into the soil and the neutron probe is inserted into the tube at various depths to determine water content at different depths in the soil profile (Burman et al., 1983). Determination of soil water content using the neutron probe is calculated from the ratio of the count taken at any depth to the mean standard count (Phene et al., 1990). However, measurements close to the soil surface are unreliable (Cole, 1989, and Gardner et al., 1991) because fast neutrons can escape to the atmosphere (Cuenca, 1989). Neutron probes have been widely used for measuring soil moisture content, but the radioactive source requires careful handling and is concern to some potential users. Time Domain Reflectometry (TDR). Time domain reflectometry (TDR) is used to measure soil water content based on the propagation of an electric signal which is dependent on the dielectric constant of the soil (Phene et al., 1990; Paltineanu and Starr,

49 1997). Parallel transmission lines are installed in the soil and serve as conductors, while the soil serves as the dielectric medium between them (Phene et al., 1990). A pair of rods acts as wave-guides and the signal propagates as a plane wave in the soil (Topp and Davis, 1985). A high-frequency signal is delivered from the TDR receiver to the transmission lines. The signal is reflected from the end of a transmission line (Phene et al., 1990; Paltineanu and Starr, 1997) back to the TDR receiver. A timing device measures the time between sending and receiving the reflected signal. This time interval relates directly to the velocity of the signal in the soil (Topp and Davis, 1985). The velocity and amplitude of the reflected signal is used to calculate the dielectric constant. There is a strong relationship between the dielectric constant and volumetric soil water content (θV) (Topp and Davis, 1985). Topp et al. (1980) stated that the determination of θV by TDR is accurate without having to calibrate the system in each soil type. On the contrary, Werkhoven (1993) suggested calibrating TDR against volumetric soil water content for each soil in order to get accurate determinations of soil water content. Time domain reflectometry is useful in studies of real-time soil water content dynamics (Platineanu and Starr, 1997). However, this technique works well only in small areas due to the limitation of cable length, which should not be more than 25 m (Platineanu and Starr, 1997) in order not to interfere with the reflected signal. Capacitance probes. Multisensor capacitance probes measure soil moisture content based on the dielectric constant of the soil (Phene et al., 1990; Paltineanu and Starr, 1997; Wu, 1998). The dielectric constant of the soil is composed of the dielectric constants of water (80.4), soil particles (3-7) and air (1) (Robinson and Dean, 1993; Paltineanu and Starr, 1997; Starr and Paltineanu, 1998a, 1998b; Wu, 1998). Since the

50 dielectric constant of the soil particles and air are small and relatively constant compared to that of water, changes in the dielectric constant of the soil are primarily a measure of the change in soil water content. A commercially available capacitance probe system called the EnviroSCAN (Sentek PTY Ltd., Kent Town, Australia) is composed of multisensor capacitance probes with capacitance sensors, a data-logger and an internal power supply charged by a solar panel or by an external battery power (Paltineanu and Starr, 1997). The probes are inserted in a PVC tube at the desirable depth for measuring soil water content and then connected to data-logger via a cable (Paltineanu and Starr, 1997). Prior to installation, the data-logger is configured to the number of probes and sensors that will be installed, and normalized for air and water counts by inserting the probe in a tube surrounded by water. The system can be set up to record the sensor reading in intervals, ranging from 1 to 9,999 min and provides real-time measurements of the change in volumetric soil water content. The data can be downloaded to a computer (Paltineanu and Starr, 1997; Starr and Paltineanu, 1998a, 1998b; Mead et al., 1994) and sensor readings can be converted to volumetric soil water content using calibration equations stored in the data-logger (Paltineanu and Starr, 1997). The system comes with software that is designed for irrigation scheduling. Conclusions The goal of irrigation scheduling of fruit crops is to meet tree water requirements for optimum yield by determining the water quantity and frequency of irrigation. Lack of irrigation scheduling may lead to excessive or inadequate water application. Excessive irrigation can result in waterlogging, potential chemical leaching from the root zone into groundwater, and increased production costs. On the other hand, a poor estimation of tree water requirements may result in water stress resulting in adverse effects on tree

51 physiological processes, water relations, vegetative growth, flower and fruit production and quality. Measurements of soil water content, plant water status or climatic data are all practical and feasible methods of irrigation scheduling of fruit crops. Irrigation scheduling based on soil water content has been widely adopted due to the availability and ease of use of various devices that monitor soil water status either discretely (e.g. tensiometers, neutron probes) or continuously (e.g. multisensor capacitance probes). Irrigation scheduling based on tree water status has gained interest in recent decades. Several studies have determined that tree stem water potential, gas exchange measurements, sap flow, and stem daily shrinkage, among other tree measurements, can be utilized for making decisions about irrigation of fruit crops. For several fruit crops, yield is more highly correlated with ΨS than with soil water potential, thus validating the use of basing irrigation decisions on plant water status rather than soil water content. There have been a few studies comparing soil water content to plant water physiological variables as bases for irrigation scheduling. Carambola is cultivated in many tropical and subtropical regions of the word. It has among the highest economic returns of any fruit crop in southern Florida. The tree is well adapted to Krome very gravelly loam soils, and produces two crops per year. Tree water requirements have not been determined; neither has the long-term response to various levels of soil moisture under subtropical climatic conditions of South Florida. Therefore, carambola can serve as a good model crop for comparing soil water content to plant water status determinations for making irrigation decisions for subtropical fruit tree species. Determination of soil water content can be achieved by evaluating and

52 calibrating various devices for use in Krome soils, as well as determining the impact of various soil moisture levels on carambola tree phenology, physiology, growth, yield, and fruit quality.

CHAPTER 3 CALIBRATION OF THE NEUTRON PROBE AND MULTISENSOR CAPACITANCE PROBES IN A GRAVELLY LOAM CALCAREOUS SOIL There is a lack of quantitative information about water requirements of tropical fruit crops in calcareous soils. Several methods and devices have been tested in these soils, including, tensiometers, neutron thermalization probes, and capacitance probes. Gravimetric determination of water content is generally not practical for irrigation scheduling in these very hard soils because sampling at different depths in the soil profile is labor intensive, time consuming, and requires special digging tools. In calcareous soils in South Florida, tensiometers (Richards, 1942) have been used for irrigation management of vegetable (Li et al., 1998) and fruit crops (Núñez-Elisea et al, 2000) due to their low cost and simple installation. However, because of the gravelly texture of soil, tensiometers were not effective with tension greater than 20 kPa due to discharge from the water column or air entering into the suction cup (Al-Yahyai et al., 2003; NúñezElisea et al, 2001). Fruit trees can tolerate higher soil tensions than herbaceous annual crops where irrigation at soil tensions less than 20 kPa are essential to avoid water stress and reduction in growth and yield (Li et al., 1998). Therefore, tensiometers may not be practical tools for irrigation scheduling of fruit crops in oolitic limestone soils. Monitoring soil water content using a neutron probe is a reliable method of irrigation scheduling for fruit crops. Monitoring soil water content based on the dielectric constant of the soil has long been used in time domain reflectometry (TDR) (Topp et al., 1980). Technology utilizing

53

54 this concept is employed in multisensor capacitance probes, whereby sensors installed at various depths in the soil profile can measure soil water content continuously (Paltineanu and Starr, 1997; Phene et al., 1990; Wu, 1998). A multisensor capacitance probe system (EnviroSCAN, Sentek PTY Ltd., Kent Town, Australia) was found useful for irrigation scheduling of lime, avocado and carambola trees (Zekri et al., 1999; Núñez-Elisea et al, 2001) in calcareous soils in South Florida. Due to the variability in soil properties, calibration equations for each soil type are essential to accurately assess soil water content (Baumhardt et al., 2000; Hanson and Peters, 2000; Morgan et al., 1999; Paltineanu and Starr, 1997). Depending on the nature of the soil, calibration of the EnviroSCAN system can be carried out in the field (Morgan et al., 1999) or in containers (Paltineanu and Starr, 1997). A calibration curve has been developed for the fine sandy soils of Florida (Morgan et al., 1999), but the system has not been calibrated for use in oolitic limestone soils in the tropical fruit production areas of southern Florida. The relationship between soil water tension as measured with tensiometers and volumetric water content is known as a soil water characteristic curve or soil water retention curve. This curve changes with soil texture and it is unique to each soil. Soil water retention curves for a calcareous soil in South Florida have been described by Muñoz-Carpena et al. (2002) and Núñez-Elisea et al. (2001) in an orchard using capacitance probes to directly measure volumetric water content. Núñez-Elisea et al. (2001) found that a soil water retention curve was better correlated with soil water tension at 10 than at 30 cm due to the difference in soil texture between the two depths. In a carambola orchard in a calcareous soil in South Florida, Al-Yahyai et al. (2003) observed that tensiometer readings did not reflect volumetric soil water content at a 10-

55 cm depth when soil water tension reached greater than 10 kPa. The large changes in tension that correspond to small changes in soil water content is a characteristic soil water retention pattern of rock-plowed calcareous soils of South Florida (Muñoz-Carpena et al., 2002). The objective of this study was to calibrate a neutron probe and a multisensor capacitance probe system (EnviroSCAN) in Krome very gravelly loam soil, a calcareous soil classified as a loamy-skeletal, carbonatic, hyperthermic, Lithic Udorthents (Noble at al., 1996). Materials and Methods Capacitance probe calibration. Calibration of multisensor capacitance probes was conducted in 15 L containers of Krome gravelly loam soil [loamy-skeletal, carbonatic, hyperthermic, Lithic Udorthents (Noble et al., 1996)] and placed above ground on a perforated plastic pad in Homestead, Fla. The soil was obtained from a location adjacent to a tropical fruit orchard, oven dried at 105 oC for 5 d and then placed in the containers. The soil was packed every 5 cm as it was added to the container to maintain equal bulk density of approximately 1.4 g·cm-3 throughout the soil profile to a total volume of 11.44 L. One probe was installed in a polyvinyl chloride (PVC) access tube. The capacitance sensor was centered at 5 cm below the soil surface. The radius of the soil in the containers was approximately the same radius of the soil water detection sphere of the capacitance sensor (10 cm). At the beginning of the experiment, the containers were placed inside larger containers and gradually flooded from the bottom to displace air from the soil. The soil was saturated for 3 h and then the containers were weighed every 2 h, three times on the first day and twice a day (at 1000 and 1600 HRS) thereafter for 13 d. The capacitance probe system was set to record data every 10 min for the duration of the experiment.

56 Gravimetric soil water content was determined by weighing the containers. Volumetric water content (θv) was calculated as the percentage of depleted soil water per time interval divided by the soil volume of the container. The calibration equation for the multisensor capacitance probe was obtained from the relationship between volumetric soil water content and capacitance probe scaled frequency in order to obtain the coefficients for the system to correctly calculate the volumetric soil water content. The scaled frequency was calculated from the following equation [SF = (Fa-Fs)/(Fa-Fw)], where SF is the scaled frequency of oscillation of the EnviroSCAN capacitance sensor, Fs is the sensor frequency reading in the soil, Fa is the sensor frequency reading in the air, and Fw is the sensor frequency reading in the water (Paltineanu and Starr, 1997). Neutron probe calibration. A neutron probe (Model 503DR, Campbell Pacific Nuclear, Inc., Martinez, Calif.) was calibrated in 95 L containers in Krome very gravelly loam soil. The soil volume in the container was within the soil water detection sphere of the neutron probe. The radius (cm) of the detection sphere was 21.4 cm as calculated with the Kristensen (1973) equation (R = 100 / (1.4 + 0.1W), where W is the volumetric water content which was 32.71%. One PVC access tube was placed in the center of each of the three containers for neutron probe measurements. The containers were filled with Krome very gravelly loam soil that was collected from a site adjacent to a tropical fruit orchard in Homestead, Fla. Prior to filling the containers, the soil was oven dried at 105 oC for 5 d. The soil was packed from the bottom of the container every 10 cm to maintain a bulk density of about 1.4 g·cm3 throughout the soil profile. The total soil volume was 88.45 L per container. The containers were placed above ground on a perforated plastic pad. The bottom of the containers was flat with multiple holes to allow free water drainage. The

57 containers were filled with water and the soil was allowed to become saturated overnight. Prior to installing the probe into the soil, 10 standard counts were taken 1 m above the soil surface. Raw counts were recorded during 32-s intervals at a depth of 20 cm once a day (at 1600 HR) and the containers were weighed simultaneously with a heavy-duty scale (Model CD-11, OHAUS Corp., Florham Park, N.J.). The calibration equation for the neutron probe was obtained from the relationship between the soil water content and the neutron probe count ratio. Count ratios were calculated by dividing the raw counts taken at a 20-cm depth by the average standard count taken at 1 m above ground level while the probe was in the shield (Hanson and Dickey, 1993). Field soil water retention curves. Soil water retention curves were determined in an 8-year-old carambola orchard in Krome very gravelly loam soil in Homestead, Fla. Soil matric potential was determined with using low-tension tensiometers (0 to 40 kPa) (Model LT, Irrometer Co., Inc., Riverside, Calif.), and soil volumetric water content measured with multisensor capacitance probes (EnviroSCAN, Sentek PTY Ltd., Kent Town, Australia) and a neutron probe (Model 503DR, Campbell Pacific Nuclear, Inc., Martinez, Calif.). Tensiometers were calibrated prior to installation using a vacuum chamber (Smajstrla and Pitts, 1997) to ensure that the water column in the tensiometer was free of air, that there were no leaks, and to standardize gauge readings among tensiometers. Three tensiometers were installed 60 cm from the trunk of each of four trees at 10, 20 and 30 cm depths. Prior to installation, a hole was made in the soil, slightly larger than the diameter of the tensiometer. A slurry prepared with sieved Krome very gravelly loam soil mixed with water was poured into the tensiometer hole to ensure that the ceramic cup was in contact

58 with the soil (Núñez-Elisea et al., 2001). Tensiometers were regularly maintained in the field using a hand-pump and water was refilled whenever it drained from the tensiometer tube. One 85-cm long polyvinyl chloride (PVC) access tube was installed at 60 cm from the trunk of each of the four replicate trees. The neutron probe was placed in the access tubes and lowered to 10, 20, and 30 cm depths. Neutron probe readings were recorded for 32 s at each depth. Multisensor capacitance probes were installed in the orchard following the procedure described by Paltineanu and Starr (1997). One probe was installed inside a 74 cm-long PVC access tube at 60 cm from the trunk of each of four replicate trees. Three sensors were placed in each probe at 10-, 20-, and 30-cm depths. The PVC tubes were installed with a motorized drill and slurry was then added to the hole. The slurry consisted of 2:1:1 by volume of calcareous rock, cement, and water to prevent air gaps from forming between the tubes and the surrounding soil (Núñez-Elisea et al., 2001). The sensors in each probe were connected to a data-logger powered by a 12-volt battery charged by a solar panel. Data were recorded every 30 min and downloaded from the data-logger to a portable laptop computer where graphs of soil water depletion rates at each soil depth and location were created with EnviroSCAN software. Soil water retention curves were calculated based on van Genuchten’s model (1980): θ(Ψ) = θr + (θs – θr) [1 + (α|Ψ|n)]-m

59 Where Ψ is the soil matric potential, θr is the residual water content, θs is the saturated water content, and α, n, and m are fitting parameters directly dependent on the shape of the θ(Ψ) curve. Results and Discussion Capacitance probe calibration. The relationship between capacitance probe scaled frequency (SF) and volumetric soil water content (θv) was linear (SF = 0.011 θv + 0.5206; r2 = 0.98) (Fig. 3-1). The coefficients from this equation were A = 0.011, B = 1, C = 0.5206. These coefficients can be used instead of the default EnviroSCAN equation for an accurate determination of volumetric water content in Krome soil. Volumetric soil water content (θv) is calculated by the EnviroSCAN based on the following equation (Paltineanu and Starr, 1997):
⎛ SF − C ⎞ θv = ⎜ ⎟ ⎝ A ⎠
1 B

, where θv is volumetric water content and A, B, and C are the

coefficients determined above. Neutron probe calibration. The relationship between volumetric soil water content and neutron probe count ratio (x) was linear (θv = 29.05 x - 6.4; r2 = 0.95) (Fig. 32). Calibration relationships of neutron probes are generally linear through the common range of soil moisture (Merriam and Knoerr, 1961). Field soil water retention curves. In Krome soil, soil water tension measured with a tensiometer was more closely correlated with capacitance probe measurements of volumetric water content at a depth of 10 cm (r2 = 0.86) than at 20 cm (r2 = 0.62 ), with the lowest coefficient of determination at 30 cm (r2 = 0.38) (Fig. 3-3, Table 3-1). This is in agreement with data obtained for Krome soil by Núñez-Elisea et al. (2001), who found that the volumetric soil water content determined with multisensor capacitance probes

60 was less strongly correlated with soil water tension at a 30-cm than at 10-cm depth due to the larger percentage of gravel at the 30-cm depth. Soil water retention curves for Krome soil can be divided into two tension ranges: from 0-10 kPa for gravel and 10-40 kPa for loam soils, where volumetric water content did not correspond to a change in soil tension (Muñoz-Carpena et al., 2002). Soil water retention curves were obtained from fitting volumetric soil water content measurements using a neutron probe with tensiometer readings based on van Genuchten’s model (1980) (Fig. 3-4). Coefficients of determination between observed and fitted data of the soil water retention curves were higher than those obtained from the capacitance probes. Coefficients of determination at different depths in the soil profile were 0.94 at 10 cm, 0.68 at 20 cm, and 0.88 at 30 cm (Table 3-2). Volumetric soil water content measurements obtained with the neutron probe were more closely correlated to soil water tension than those obtained by capacitance probes. The capacitance sensors may have been less accurate than the neutron probe due to an increased amount of air from coarse textured soil which was greater at lower depth in the soil profile (NúñezElisea et al., 2001). Differences in precision of the neutron probe and capacitance probes can be caused by differences in interference, measurement methods, and sphere area of the soil water detection zone (Paltineanu and Starr, 1997). Nonetheless, several studies suggested that the neutron probe was more consistent than capacitance probes under field conditions (Evett, 2000; Evett and Steiner, 1995; Evett et al., 2002; Heng et al., 2002). This is especially true in coarse textured soil such as Krome very gravelly loam where the potential for air gaps is high (Al-Yahyai et al., 2003; Tomer and Anderson, 1995).

61 Conclusions The calibration equation obtained for the capacitance probe was y = 0.011x + 0.5206; r2=0.98, and that for the neutron probe was y = 29.05x-6.4; r2=0.95. In Krome very gravelly loam soil, soil water retention curves of tension vs. volumetric water content using capacitance probes or a neutron probe were fitted with van Genuchten’s model. The r2 values indicated that tensiometers are more highly correlated with neutron probe measurements of volumetric soil water content than with multisensor capacitance probe measurements. This suggests that capacitance probe measurements of soil water content were more variable in response to changes in soil texture throughout the soil profile.

62

Table 3-1. Fitted parameters of van Genuchten’s model used to describe soil water retention curves of Krome very gravelly loam soil where soil water content was measured in a carambola orchard using multisensor capacitance probes (EnviroSCAN) at soil depths of 10, 20, and 30 cm. Parametersy 10 cm 20 cm 30 cm θs θr α n m r2 z
z 2 y

0.47 0.20 0.72 1.56 0.36 0.86

0.47 0.29 0.87 1.98 0.50 0.62

0.47 0.28 0.10 22.51 0.96 0.38

r values for observed vs. fitted values, n = 4. θs is saturated water content, θr is residual water content, and α, n, and m, are model fitting parameters.

63 Table 3-2. Fitted parameters of van Genuchten’s model used to describe soil water retention curves of Krome very gravelly loam soil where soil water content was measured in a carambola orchard using a neutron probe at soil depths of 10, 20, and 30 cm. Parametersy 10 cm 20 cm 30 cm θs θr α n m r
y 2z z 2

0.47 0.06 0.44 1.22 0.18 0.94

0.47 0.19 4.10 1.22 0.18 0.69

0.470 0.08 4.63 1.14 0.13 0.88

r values for observed vs. fitted values, n = 4. θs is saturated water content, θr is residual water content, and α, n, and m, are model fitting parameters.

64

0.78 0.76 0.74

Scaled Frequency (SF)

0.72 0.70 0.68 0.66 0.64 0.62 0.60 0.58 0.56 6 8 10 12 14 16 18 20 22 24

θv (%)

Fig. 3-1. Relationship between scaled frequency (SF) from a multisensor capacitance probe (EnviroSCAN) and volumetric water content (%) fitted to obtain calibration coefficients for Krome very gravelly loam soil in containers, y = 0.011x + 0.5206; r2 = 0.98.

65

34

32

30

θv (%)

28

26

24

22

20 0.9 1.0 1.1 1.2 1.3 1.4

Count Ratio

Fig. 3-2. Relationship between count ratio from the neutron probe and volumetric water content (%) fitted to obtain a calibration equation for Krome very gravelly loam soil in containers, y = 29.05x - 6.4; r2 = 0.95.

66

40

10 cm r2 = 86

30

θv (%)
20 10 0

40 T(cm) vs ES-10 h=(T*10) vs ES-10-k 30

20 cm r2 = 62

θv (%)
20 10 0

40

30 cm r2 = 38

30

θv (%)
20 10 0 0 20 40 60 80 100 120 140

Soil water tension (cm)

Fig. 3-3. Soil water retention curves for Krome very gravelly loam soil at 10-, 20-, and 30-cm depths based on van Genuchten’s model calculated from volumetric water content (θv) measured with capacitance probes and soil water tension measured with tensiometers.

67

10 cm r2 = 94
40

30

θv (%)
20 10 0

20 cm r2 = 68
40

30

θv (%)
20 10 0

30 cm r2 = 88
40

30

θv (%)
20 10 0 0 20 40 60 80 100 120 140

Soil water tension (cm)

Fig. 3-4. Soil water retention curves for Krome very gravelly loam soil at 10-, 20-, and 30-cm depths based on van Genuchten’s model calculated from volumetric water content (θv) measured with a neutron probe and soil water tension measured with tensiometers.

CHAPTER 4 MONITORING SOIL WATER CONTENT FOR IRRIGATION SCHEDULING IN A CARAMBOLA ORCHARD IN A GRAVELLY LIMESTONE SOIL There are approximately 100 ha of carambola trees in Florida (J. H. Crane, University of Florida, personal communication), of which 46 ha are in Miami-Dade County (Degner et al., 2002). A sweet-type, ‘Arkin’, is the leading commercial carambola cultivar in Florida (Knight and Crane, 2002). Excessive soil water content (Joyner and Schaffer, 1989) and drought (Ismail and Noor, 1996a; Ismail et al., 1996; Salakpetch et al., 1990) decrease carambola growth and yield. In South Florida, 86% of the total rainfall in 2001 and 79% of the total rainfall in 2002 occurred between May and October (Fig. 4-1). During the winter and drought periods, irrigation may be needed to compensate for the lack of water from precipitation. Therefore, irrigation based on quantitative information is important for commercial carambola production in south Florida. The soil of the Miami-Dade County, where carambola is cultivated, is composed primarily of calcium carbonate (Degner et al., 1997) and classified as Krome very gravelly loam. This is a very shallow, mineral soil with a pH of 7.4 - 8.4 (Noble et al., 1996). This soil is low in organic matter and commercial farming largely depends on fertilizer applications (Degner et al., 2002). The high demand for fertilizer coupled with potentially excessive irrigation creates a potential for agrochemical leaching into the groundwater (Muñoz-Carpena et al., 2002; Zekri et al., 1999). In addition to reducing potential agrochemical leaching, scheduling irrigation to apply only the amount of water

68

69 required by the plant should increase grower economic returns by reducing fertilizer and water inputs, and improving plant growth and yields. A survey by Li et al. (2000) in 1998 showed that 73% of tropical fruit growers in Miami-Dade County schedule irrigation based on the frequency and quantity of rain. The percentage declined to 64.3% in 2002 according to a more recent survey by MuñozCarpena et al. (2003). The number of growers monitoring of soil water content for irrigation scheduling has increased to 48.8% of the respondents to the 2002 water-use survey (Muñoz-Carpena et al., 2003) compared to only 15% in 1998 (Li et al., 2000). According to the 1998 survey, methods of determining soil water included tensiometers, capacitance probes, digging and squeezing soil, and the feel and appearance of the soil (Li et al., 2000). Duration and frequency of irrigation were highly variable among tropical fruit growers, which highlight the need for a better understanding of irrigation requirements of these crops. With overhead, high-volume sprinklers, irrigation was applied from one to three times per week for one to 12 h per application (Li et al., 2000). For microsprinklers, the frequency of operation ranged from 0.5 to 7.5 h per application and from one to seven per week. The amount of water applied per tree ranged from 110 to 2302 L with overhead sprinklers, 19 to 341 L per tree with microsprinklers, and from 7.6 to 45 L per tree for drip irrigation (Li et al., 2000). The variability in responses to irrigation quantities and frequencies used by growers can be attributed to a lack of basic quantitative information. Tensiometers, neutron probes, and capacitance probes monitor soil water content and are often used for irrigation scheduling. Tensiometers measure soil suction or matric water potential rather than soil water content (Richards, 1942; Smajstrla and Harrison,

70 1998). A soil water retention curve must be established to determine the soil water content that corresponds to the water matric potential in order to estimate soil water content with a tensiometer. The neutron probe is a portable compact unit that is easy to operate and volumetric soil water content can be obtained instantly at different depths of the soil profile. Neutron probes are considered more accurate than other soil water content monitoring devices for irrigation scheduling (Evett and Steiner, 1995; Mostert and Hoffman, 1996). The main component of a neutron probe is a fast neutron source encased in a protective shield and an electronic counting scaler which are connected by an electric cable that is also used to lower the probe into an access tube to determine soil water content throughout the soil profile (Chanasyk and Naeth, 1996). Neutrons with a high energy are emitted by a radioactive source, such as americium 241/beryllium, into the soil and are slowed down by collisions with nuclei, primarily hydrogen atoms (Gardner and Kirkham, 1952). The density of the resultant cloud of slow neutrons is a function of the soil water content (Chanasyk and Naeth, 1996). The number of fast neutrons that are slowed is detected and measured as a ‘count rate’ per unit time (Gardner et al., 1991). The count rate is converted to volumetric water content using a calibration curve. Neutron probes measure soil volumetric water content as the percentage of water per volume of soil. Capacitance probes measure the soil water content based on the dielectric constant of the soil (Paltineanu and Starr, 1997; Phene et al., 1990; Wu, 1998) a concept that was first proposed for soil monitoring by time domain reflectometry (TDR) (Topp et al., 1980). The dielectric constant of the soil is composed of the dielectric constants of water (80.4), soil particles (3-7) and air (1) (Robinson and Dean, 1993; Paltineanu and Starr,

71 1997; Starr and Paltineanu, 1998a; Wu, 1998). The dielectric constant of the soil particles and air are small and relatively constant compared to that of water. Thus changes in the dielectric constant of the soil are a measure of the change in soil water content. The volumetric water content can be expressed either as a percentage or a depth of water (mm of water / 10 cm of soil) (Núñez-Elisea et al., 2001; Paltineanu and Starr, 1997; Starr and Paltineanu, 1998a). The objective of this study was to evaluate and compare tensiometers, multisensor capacitance probes, and a neutron probe for accurately assessing soil water content in a carambola orchard in Krome very gravelly loam soil. Materials and Methods The experiment was conducted in an orchard of 8-year-old ‘Arkin’ carambola trees grafted onto open-pollinated Golden Star rootstock at the Tropical Research and Education Center in Homestead, Fla. Trees were spaced at 4.5 m within rows and 6.1 m between rows. Low-tension tensiometers (0 to 40 kPa) (Model LT; Irrometer Co., Inc., Riverside, Calif.) were calibrated prior to installation using a calibration vacuum chamber (Smajstrla and Pitts, 1997) to ensure that the water column in the tensiometer was air free and that there were no leaks. One tensiometer was installed 60 cm from the trunk of each of the three replicate trees in each treatment. Tensiometers were installed at a depth of 10 cm. Prior to installation, a hole was made in the soil, slightly larger in diameter than the tensiometer. A slurry, prepared with sieved soil mixed with water, was applied to the tensiometer hole to ensure that the ceramic cup of the tensiometer was in contact with the soil (Núñez-Elisea et al., 2001). Tensiometers were maintained regularly in the field using a hand-pump and water was refilled whenever it drained from the tensiometer tube.

72 For neutron probe measurements, one 85-cm long polyvinyl chloride (PVC) access tube was installed at 60 cm from the trunk of each of three replicate trees in each of the four treatments. The neutron probe (Model 503DR, Campbell Pacific Nuclear, Inc. Martinez, Calif.) was placed in the access tubes and lowered to 10, 20, 30, and 50 cm depths below the soil surface. Neutron probe counts were recorded during 16-s intervals were obtained at each depth. A multisensor capacitance probe system (EnviroSCAN, Sentek PTY Ltd., Kent Town, Australia) was used to automatically and continuously measure soil water content in each treatment. Prior to installation, dataloggers were configured in the laboratory following the procedure described by Paltineanu and Starr (1997). Sensors were normalized to air and water counts by placing the probe in a tube surrounded by water. One probe was installed inside a 74 cm-long PVC access tube at 60 cm from the trunk of each of the three replicate trees in each of the four treatments. Four sensors were placed in each probe at 10, 20, 30, and 50 cm below the soil surface. The PVC tubes were installed with a motorized drill and slurry was then poured into the hole. The slurry consisted of 2:1:1 by volume of calcareous rock, cement, and water to prevent air gaps from forming between the tubes and the surrounding soil (Núñez-Elisea et al., 2001). The sensors in each probe were connected to a datalogger powered by a 12-volt battery charged with a solar panel. Data were recorded every 30 min and downloaded from the datalogger to a portable laptop computer and graphs of soil water depletion rates at each soil depth and location were created with the EnviroScan software. Trees were separated into four irrigation treatments based on the rate of soil water depletion (SWD) between field capacity and the first visual sign of stress (i.e., leaf

73 yellowing and abscission) of carambola trees (as measured with multisensor capacitance probes in a preliminary experiment). The SWD levels were: 0-8% SWD, 9-11% SWD, 12-14% SWD, or 15-17% SWD. Treatments were arranged in a completely randomized design with three replications per treatment. When the soil water content reached the treatment range, trees were irrigated with microsprinklers (at 89 L·h-1) to restore the soil to field capacity. Data from the three instruments were compared and analyzed by linear and nonlinear regression and correlation tests. Results and Discussion A soil water retention curve was developed using van Genuchten’a (1980) model: θ(Ψ)= θr + (θs – θr) [1+(α|Ψ|)n]-m where Ψ is the matric potential (suction or water tension), θr is the residual water content, and θs is the saturated water content, and α, n, and m are fitting parameters directly dependent on the shape of the θ(Ψ) curve. Parameters (i.e. θr, θs, α, n, and m) for van Genuchten’s model for the soil water retention curve of Krome soil were obtained from the relationship between soil matric potential (Ψ) measured by tensiometers and volumetric water content (θ) determined by neutron and capacitance probes (Table 4-1). Soil matric potential measured with tensiometer fit van Genuchten’s model better when volumetric soil water content was measured with a neutron probe (r2 = 0.42) than with multisensor capacitance probes (r2 = 0.35) (Fig. 4-2 and 4-3, respectively). The fairly weak relationship between soil water tension and volumetric soil water content in Krome very gravelly loam soils can be attributed to the inaccuracy of tensiometer readings above a suction of 20 kPa and heterogeneity of very gravelly Krome soils (Núñez-Elisea et al., 2001). Tensiometers installed at 10 cm depth were not effective at a tension of above 20

74 kPa because air entered into the suction cup through the large pores in the gravel and produced inaccurate measurements or water discharged completely from the tensiometer. In a previous study, tensiometers installed at a 10-cm depth in Krome very gravelly loam soil in south Florida were successfully used to schedule irrigation of tomato (Li et al., 1998). In contrast, a previous study in tropical fruit orchards with the same soil showed that tensiometers were not effective in accurately estimating soil water potential at a depth of 30 cm (Núñez-Elisea et al., 2001). The difference between the usefulness of tensiometers in the vegetable field and fruit orchard may have been a result of significantly larger soil particles in the fruit orchards. In south Florida, vegetable fields are rock-plowed and repeatedly disked to break up the top layer of the soil (Colburn and Goldweber, 1961). Núñez-Elisea et al. (2001) reported that the lack of effectiveness of tensiometers in tropical fruit orchards at a depth of 30 cm was attributed to the rockiness of Krome soil at that depth where 71 to 73% of the soil was gravel compared to 26% to 38% at in top 10 cm. Similar relationship between tensiometer readings and capacitance probes readings were also observed in the present study. Muñoz-Carpena et al. (2002) measured gravimetric water content and soil suction in a laboratory experiment. They reported that at soil suction above 10 kPa, soil water content in Krome very gravelly loam soil is relatively insensitive to tension changes, thus large changes in soil water tension reflect small changes in actual soil water content. In the present study, soil suction above 10 kPa resulted in variable tensiometer readings and discharge from the water column above 20 kPa. Thus, tensiometers are not very useful for monitoring soil water content for irrigation scheduling in carambola orchards in Krome very gravelly loam soil.

75 Volumetric soil water content determined with a neutron probe and capacitance probes were positively correlated at all depths (Fig. 4-4). However, the correlation was not very high, probably due to differences in the principles of operation of the devices, variability in soil microclimate around the access tubes, and the larger sphere of influence (volume of soil measured by the probe) measured by the neutron probe than each capacitance sensor. Neutron probe measurements close to the soil surface are not reliable (Chanasyk and Naeth, 1996; Cole, 1989; Gardner, et al., 1991;) due to escape of fast neutrons to the atmosphere (Cuenca, 1989) as a result of the radius of the sphere of influence of neutron probes (approximately 20 cm; Chanasyk and Naeth, 1988). This explains the overestimation of soil water content observed in this study (Fig. 4-2) compared to values estimated by Muñoz-Carpena et al. (2002) for the same soil using pressure plate apparatus in the laboratory. Of the three soil water monitoring devices we tested, variability among readings at different locations at the same depth were the lowest with the neutron probe [coefficient of variation (CV) = 11%] compared to tensiometers (CV = 17%) or capacitance probes (CV =33%). Although neutron probe readings were less variable than those obtained from capacitance probes and tensiometers, its use is less practical than that of the other two devices for irrigation scheduling due to the radioactive source of fast neutrons which can cause health hazards if the neutron probe is not used properly. Moreover, neutron probe operation and transportation require permits and continuous monitoring for radiation leakage. Specific calibration is required for accurately assessing actual volumetric soil water content. The use of a neutron probe to monitor soil water content in Krome soil is labor intensive because it requires installation of several access tubes using a motorized-drill and the manually obtaining of readings

76 over a large area at different depths in the soil profile. The initial cost of the instrument is very high compared to that of tensiometers. Studies have shown that volumetric soil water content determined with multisensor capacitance probes was highly correlated with volumetric soil water content determined gravimetrically (Paltineanu and Starr, 1997) in Mattapex silt loam fine soil, in silty clay loam soils (Ould Mohamed et al., 1997), in fine sand soils of Florida (Morgan et al., 1999), and in weathered heterogeneous soils (Wu, 1998). Multisensor capacitance probes accurately measured soil water content for citrus orchards in Candler fine sand (Fares and Alva, 2000). They also proved reliable for irrigation scheduling in avocado and ‘Tahiti’ lime orchards in south Florida (Núñez-Elisea et al., 2001; Núñez-Elisea et al., 2000; Zekri et al., 1999). However, in the present study, there were often variable readings among sensors at the same depth receiving the same irrigation treatment. Inaccurate measurements of volumetric water content by capacitance probes were reported by Hanson and Peters (2000) in silty loam and silty clay soils in California, and by Evett and Steiner (1995) in Amarillo fine sandy loam soil. Tomer and Anderson (1995) concluded that for coarse textured soils, changes in water content are difficult to detect using capacitance probes. Although not proven, we speculate that increased capacitance probe variability in Krome soil over time could be attributed to temperature variations from the winter to the summer, soil wetness and drying cycles, long-term changes in soil chemical and physical properties, and the initial soil disturbance caused by trenching. In addition, air gaps and disturbance of soil around the probe may lead to a change in soil bulk density that can produce measurement errors (Paltineanu and Starr, 1997). Throughout this study, lightning had a devastating effect on electrical connections and capacitance

77 sensors even when the system was surge protected and a grounding rod was installed. At times, capacitance sensors briefly produced erroneous readings, such as sudden decreases in soil water readings, spikes, or discontinuous readings. However, the system generally recovered from these temporary disturbances. Variability in capacitance probe readings in this study can also be attributed to a low battery or a damaged solar panel and condensation of water vapor in the probes. To reduce water vapor condensation in access tubes, silica gel desiccants were frequently replaced but due to the large tube size, this may not have been sufficient to maintain a dry climate inside each probe. A major advantage of irrigation scheduling using capacitance probes is that scheduling is based on the rate of soil water depletion rather than on the absolute water content. Therefore, capacitance probe measurements reflect the change in soil water content and evapotranspiration over time (Zekri et al., 1999). Using Sentek EnviroSCAN software, the data are presented graphically and options are provided for the user to divide the display into separate zones to manage soil water content. Multisensor capacitance probe system can be a useful tool for irrigation scheduling if the cost of operation and maintenance is financially justified. In addition to tensiometers, neutron probes, and multisensor capacitance probes, there are several other methods of monitoring soil water status that include dielectric sensors, volumetric water content measurement devices, and water suction devices. Several of these devices are currently being tested for use in Krome very gravelly loam soils at the Tropical Research and Education Center in Homestead, Fla.

78 Table 4-1. Fitted parameters of van Genuchten’s model used to create a soil water retention curves for Krome soil. Soil water content was measured in a carambola orchard with a neutron probe and multisensor capacitance probes. Parametersz θs θr α n
z

Neutron probe 0.4758 0.226 0.029 2.67

Capacitance probes 0.4758 0.229 0.0598 2.41

0.585 0.625 m θs is saturated water content, θr is residual water content, and α, n, and m, are model fitting parameters.

79

Amount of rainfall and ET (mm)

350 300 250 200 150 100 50 0
Ja n Fe -0 1 b M -01 ar Ap -01 r M -01 ay J u -01 nJu 0 1 Au l-01 g Se -01 p O -01 ct N -01 ov D -01 ec J a -01 n Fe -0 2 b M -02 ar Ap -02 r M -02 ay J u -02 nJu 0 2 Au l-02 g Se -02 p O -02 ct N -02 ov D -02 ec -0 2

Rain ET

Fig. 4-1. Total monthly rainfall and evapotranspiration (ET) during 2001 and 2002 in Homestead, Fla. Source: Florida Automated Weather Network, IFAS, Univ. of Fla., Gainesville.

80

50
Neutron probe volumetric water content (%)

45 40 35 30 25 20 15 10 5 0 0 10 20 30 40 50

Soil water tension (kPa)

Fig. 4-2. Relationship between volumetric soil water content measured with a neutron probe and soil water tension measured with a tensiometer in a carambola orchard in Krome very gravelly loam soil. The response line was fitted using the van Genuchten equation (r2 = 0.42).

81

50
Capacitance probe volumetric water content (%)

45 40 35 30 25 20 15 10 5 0 0 10 20 30 40 50
Soil water tension (kPa)

Fig. 4-3. Relationship between volumetric soil water content measured with multisensor capacitance probes and soil water tension measured with a tensiometer in a carambola orchard in Krome very gravelly loam soil. The response line was fitted using the van Genuchten equation (r2 = 0.35).

82

45 Neutron probe soil water content (%) 40 35 30 25 20 15 15 20 25 30 35 40 Capacitance probe soil water content (%)

Fig. 4-4. Relationship between soil water content (%) as measured by multisensor capacitance probes and a neutron probe in Krome very gravelly loam soil (y = 0.67 x + 12.19, r2 = 0.35).

CHAPTER 5 PHYSIOLOGICAL RESPONSES OF CARAMBOLA TREES TO WATER DEPLETION IN KROME VERY GRAVELLY LOAM SOIL Physiological processes in fruit trees such as water potential and gas exchange are sensitive to changes in soil water content. These physiological variables, growth, and fruit production should be correlated with soil water content prior to determining the appropriate amount of water to apply to an orchard. Continuous monitoring of soil water content in tropical fruit orchards in Krome very gravelly loam soils has been done with multisensor capacitance probes (Al-Yahyai et al., 2003; Núñez-Elisea et al., 2001; Schaffer, 1998; Zekri et al., 1999). Proprietary software (EnviroSCAN 4.0, Sentek PTY Ltd., Kent Town, Australia) is available for plotting soil water depletion (SWD) over time and determining of the ‘full point’ (field capacity), the ‘refill point’ (irrigation point), and the theoretical ‘onset of water stress’, the point at which the evapotranspiration rate is significantly reduced (Schaffer, 1998; Zekri et al., 1999). Field capacity (FC) has been determined with multisensor capacitance probes (Zekri and Parsons, 1999; Fares and Alva, 2000) based on Veihmeyer and Hendrickson’s concept which defines FC as the amount of water in the soil after excess water had drained and the rate of downward movement has decreased (Hillel, 1998). Irrigation decisions using SWD measurements are based on maintaining soil water content between the ‘full point’ and the ‘refill point’ before the occurrence of the ‘onset of water stress’. This approach to irrigation may be more applicable to annual, herbaceous crops than to fruit trees. Trees can tolerate longer periods of low soil water content than herbaceous plants through

83

84 specialized long-term and short-term physiological, phenological, anatomical, and morphological responses (Ludlow, 1989), such as growth reduction, stomatal closure, and osmotic adjustment (Hsiao, 1973). Carambola (Averrhoa carambola L.), a tropical fruit tree native to S.E. Asia, responds to water deficits by the ‘avoidance’ strategy (Neuhaus, 2003). This strategy involves minimizing water loss by stomatal control, leaf movement, reducing leaf area, and leaf abscission (Ludlow, 1989). The lethal leaf water potential (ΨL) level for carambola is about -2.9 MPa and the lethal leaf relative water content (RWC) is about 56% (Ismail et al., 1994). Carambola responses to soil water deficit can be divided into four phases (Neuhaus, 2003): (1) minimizing water loss by stomatal closure, initial osmotic adjustment, and leaflet movement from a horizontal to vertical orientation; (2) increasing root:shoot ratio via leaf and fruit abscission, and increasing proline levels to increase leaf osmotic adjustment; (3) reaching a stage of irreversible damage caused by chlorophyll degradation, root growth inhibition, and further leaf and fruit shedding; and (4) tree death. Rewatering the trees during the second phase results in plant recovery within a few days (Salakpetch et al., 1990). Plant ΨL decreases when water loss via transpiration (E) is greater than water uptake (Salisbury and Ross, 1985). Leaf water potential of field and container-grown carambola trees declined with increasing water stress (Ismail and Noor, 1996a; Ismail et al., 1994, 1996). Ismail et al. (1994) found a difference in ΨL of up to -2.20 MPa between stressed and well-watered carambola trees. Salakpetch et al. (1990) found that ΨL of water-stressed carambola trees (-2.0 MPa) returned to the previous level (-0.6 MPa) upon rewatering. Net CO2 assimilation (A) of carambola decreased with increased water

85 stress. Net CO2 assimilation of 6-month-old, container-grown carambola trees was 3.13 µmol·m-2·s-1 for non-stressed trees compared to 0.24 to 1.83 µmol·m-2·s-1 for water stressed trees (Ismail et al., 1994). Net CO2 assimilation was reduced when ΨL fell below -0.80 MPa (Ismail et al., 1994; Razi et al., 1992). The decrease in A was due to reduced stomatal conductance. Ismail et al. (1994) reported that the leaf photosynthetic rate was reduced when stomatal resistance exceeded 3.5 s·cm-1. Most of the studies of the responses of carambola tree to water stress in tropical climates were conducted on young trees grown in containers in greenhouses. Little is known about the response of mature and young carambola trees to soil water depletion under field conditions especially in gravelly soils in subtropical climates such as that of South Florida. The objective of this study was to test the effect of soil water depletion on stem water potential (ΨS) and leaf gas exchange of carambola in a Krome very gravelly loam soil in South Florida. Materials and Methods Experimental site and plant material. An orchard of 8-year-old ‘Arkin’ carambola trees grafted onto open pollinated Golden Star rootstock were used for this study. The trees were planted with 4.5 m within and 6.0 m between rows with an artificial (Polypropylene ribbon shade cloth) windbreak on the northern, eastern, and western perimeters and a line of sapodilla [Manilkara zapota (L.) von Royen] trees as a windbreak on the southern perimeter of the orchard. Adjacent to the orchard trees, 2year-old ‘Arkin’ carambola trees grafted onto Golden Star rootstock were planted in 95 L containers. Six 2-year-old ‘Arkin’ carambola trees grafted on Golden Star rootstock were also planted in 95 L containers and placed in a greenhouse. The containers were filled with Krome very gravelly loam soil [loamy-skeletal, carbonatic, hyperthermic, Lithic

86 Udorthents (Noble et al., 1996)] that was excavated from a site next to the orchard. Orchard trees were grown in the same soil. Fertilizer application, pest management, and fertilization followed standard practice for South Florida’s commercial carambola production (Crane, 1994). Trees were irrigated with 89 L·h-1 microsprinklers with a 360o wetting area (Maxijet, Dundee, Fla.). In the orchard and in field containers, trees were irrigated at various times and rates to provide a range of SWD over the course of the experiments. The ranges were determined from a preliminary study (Al-Yahyai, unpublished data) and based on the SWD from soil field capacity (FC) to leaf yellowing, which was the first visual sign of water stress. For orchard trees, irrigation was withheld toward the end of the experiment for 121 d from 1 Aug. to 30 Nov. 2003 to obtain high SWD values. Trees in containers in the greenhouse were divided into two treatments: (1) soil water maintained at FC and (2) no irrigation water applied. Soil water measurements. Soil water depletion was determined by continuously measuring soil water content using multisensor capacitance probes (EnviroSCAN, Sentek PTY Ltd., Kent Town, Australia). Capacitance probes were installed 60 cm north of tree trunks in the orchard and 20 cm from tree trunks in the containers. The sensors recorded soil moisture content every 30 min at depths of 10, 20, 30 and 50 cm. The data were stored in a datalogger and later downloaded to a computer for analysis. Soil water content measurements from sensors placed at the 4 depths were summed and plotted using the EnviroSCAN proprietary system software. Installation of the system in Krome soils was previously described by Al-Yahyai et al. (2003) and Núñez-Elisea et al. (2001), and

87 technical multisensor capacitance probe specifications were described by Paltineanu and Starr (1997). Leaf gas exchange and water potential measurements. Net CO2 assimilation (A), stomatal conductance to water vapor (gs), transpiration (E), and stem water potential (ΨS) were determined periodically for trees in the orchard and in containers in the field. Net CO2 assimilation, gs, E, and ΨS for the container-grown trees in the greenhouse were determined three times per week for 6 weeks from 18 Oct. to 2 Dec. 2002. Four mature leaves per tree from the orchard and three mature leaves per tree from trees in containers in the field were randomly selected for leaf gas exchange and water potential measurements from 1100 to 1400 HRS. Net CO2 assimilation, E, and gs were determined with a portable infrared gas analyzer (CIRAS II, PPSystems Ltd., Herts, UK). Measurements were made on the terminal leaflet of fully matured, sunlit leaves at a photosynthetic photon flux (PPF) of 1000 µmol·m-2·s-1 which is above the saturation point for photosynthesis of carambola (Marler et al., 1994), and reference CO2 of 375 µmol·mol-1 at a flow rate of 200 ml·min-1. Leaf and stem water potential (ΨS) was measured between 1100 and 1400 HRS using a pressure chamber (Soil Moisture Equipment Corp., Santa Barbara, Calif.). Stem water potential was determined using a fully matured leaf enclosed in a reflective plastic bag for 1 h to suppress E and allow ΨS to equilibrate with leaf water potential (Begg and Turner, 1970; McCutchan and Shackel 1992; Shackel et al., 1997). Statistical analysis. Linear regression analysis was performed with SigmaPlot software (SYSTAT Software Inc., Richmond, Cal.) and nonlinear regression analysis was performed with TableCurve 2D software (SYSTAT Software Inc., Richmond, Cal.).

88 Results and Discussion Stem water potential of trees in containers in the greenhouse was above -1.0 MPa when SWD increased from 0% to 60% above which there was a sharp decrease in ΨS (Fig. 5-1). A similar trend was observed in field-container trees with a sharp decrease in ΨS when SWD reached 50% (Fig. 5-2). For the orchard trees, SWD never reached above 30%. Within the range of SWD from 0% to 30%, ΨS remained above -1.0 MPa and did not significantly correlate with SWD (data not shown). Therefore, within a 0 to 30% SWD range, there was no significant effect of ΨS on leaf gas exchange, presumably due to sufficient soil water content in the orchard from frequent precipitation and capillary rise from the water table which was 1-2 m below the soil surface. Similarly, in a study of mango trees in Krome soil, predawn water potential ranged from -0.3 to -0.5 MPa and there were no significant differences in A or E were observed between irrigated and nonirrigated trees (Larson et al., 1989). For 6-month-old carambola trees in containers, Ismail et al. (1994, 1996) observed that withholding water for 1, 2, or 4 weeks resulted in midday ΨL of -1.3 to -1.5 MPa, -2.0 to -2.5 MPa, or -2.9 MPa, respectively, in comparison to trees irrigated at field capacity that had a ΨL of -0.9 MPa. Similarly, Salakpetch et al. (1990) found that leaves of 2-year-old carambola trees in containers wilted when midday ΨL reached -2.0 MPa, 10 d after withholding water. Leaf water potential returned to its original level (-0.6 MPa) 2-3 d after rewatering (Salakpetch et al., 1990). The ability of carambola trees to regulate their water potential was attributed to osmotic adjustment through increased levels of proline (Ismail et al., 1994) and a reduction in gs (Ismail et al., 1994, 1996; Razi et al. 1992). Leaf gas exchange of container-grown carambola trees in the field and in a greenhouse was more closely correlated with ΨS than with SWD. For example, gs

89 relationship with SWD coefficient of determination (r2) was 0.46 (Fig. 5-3) compared to r2 = 0.54 (Fig. 5-4) with ΨS. This is in agreement with studies that found ΨS as a useful indicator of water stress and thus can be used for irrigation management (Naor, 2000; Shackel et al., 1997; Stern et al., 1998). Stem water potential effects on stomatal conductance. A reduction in ΨS below -1.0 MPa resulted in reduced gs for trees in containers in the greenhouse (Fig. 5-4) and in the field (Fig. 5-5). Similarly, Razi et al. (1992) reported a decline in gs in carambola as a consequent of a decrease in ΨL. Differences between greenhouse and field trees in the response of physiological variables to SWD were presumably due to greater climatic variability in the field than in the greenhouse. For example, vapor pressure was more variable in the field [coefficient of variation (cv) = 29%] than in the greenhouse [coefficient of variation (cv) = 17%]. Stem water potential effects on transpiration. Reductions in E were related to reductions in ΨS in carambola trees in containers in the greenhouse (Fig. 5-6) and in the field (Fig. 5-7). As SWD increased, transpiration of trees in containers in the greenhouse (Fig. 5-8) and the orchard (Fig. 5-9) decreased linearly as gs decreased. This response of carambola trees is in agreement with results from Razi et al. (1992) who found a linear relationship between ΨL and E in carambola trees. Stem water potential effect on net CO2 assimilation. Net CO2 assimilation decreased as ΨS declined below about -0.5 MPa in trees grown in containers in the greenhouse (Fig. 5-10) and in the field (Fig. 5-11). Ismail et al. (1994) found that mean A of young carambola trees in containers was 3.13 µmol·m-2·s-1 for well-watered trees compared to 0.24 to 1.83 µmol·m-2·s-1 for water-stressed trees. They concluded that a ΨL

90 lower than -0.80 MPa significantly reduced A. Similarly, Razi et al. (1992) observed that A of carambola trees was reduced when water potential fell below -0.85 MPa. These results suggest that A of carambola trees can be reduced when ΨS fall below -0.5 MPa. The reduction in A with decreasing ΨS was presumably due to a decrease in gs for trees in containers in the greenhouse (Fig. 5-12) and in the field (Fig. 5-13). Ismail et al. (1994) reported that increased stomatal resistance in young carambola trees limited CO2 uptake during water stress. Reduction in A of carambola trees in containers in the greenhouse and the field in Krome soils occurred when gs was below 50 mmol·m-2·s-1 (Figs. 5-12 and 5-13). Stomatal conductance effects on transpiration and net CO2 assimilation. The effect of gs on E (Fig. 5-8 and 5-9) and A (Fig. 5-12 and 5-13) suggested that partial stomatal closure was essential to conserve water by reducing E. This partial stomatal closure resulted in a gradual decline in A as soil water initially depleted. However, as soil water depletion increased and water stress progressed, a further decline in gs (below 50 mmol·m-2·s-1) resulted in sharp decline in A. This perhaps was an avoidance mechanism by carambola trees to conserve water by reducing E, while maintaining A during periods of occasional or short-term water stress. Thus a SWD lower than 50% in Krome soils did not affect tree gas exchange. In the orchard, the SWD never reached greater than 30% (data not shown). Thus, the range of SWD in the orchard also remained below the point at which leaf gas exchange was decreased. Stem water potential relationship to leaf water potential. It has been suggested that ΨL is a less reliable measure when making irrigation scheduling decisions for fruit trees than ΨS due to its inherent variability under field conditions (Garnier and Berger,

91 1985; McCutchan and Shackel, 1992; Naor, 2000). Enclosing leaves in a reflective, dark plastic bag for 1-2 h prior to water potential measurements will reduce E until an equilibrium is achieved between the ΨL and the ΨS (Begg and Turner, 1970), thus providing a more precise method of determining tree water status (Garnier and Berger, 1985; McCutchan and Shackel, 1992; Naor, 2000; Shackel et al., 1997). In this study, ΨL and ΨS from the same branch were measured within 2 min of each other to determine their relationship. There was a strong relationship (r2 = 0.97) between ΨL and ΨS in field and greenhouse container trees (Fig. 5-14). For orchard trees (Fig. 5-15), ΨL and ΨS were also highly correlated (r2 = 0.85) despite the narrow water potential range of -0.3 to -1.1 MPa compared to that of container trees. Jones (1997) suggested that either ΨS or ΨL may be used to assess the effect of water stress on a particular tree response that is assumed being affected by tree water status. Our results with carambola trees appeared to confirm this. Conclusions An increase in SWD resulted in reduction in ΨS, A, E, and gs of carambola trees grown in containers in Krome very gravelly loam soil. In a carambola orchard, SWD never reached a point where there was a decrease in ΨS, A, E, or gs. Correlations between physiological variables were higher for container-grown trees in a greenhouse than in the field, which may have been due to greater fluctuations in vapor pressure in the field. In the orchard, SWD did reach greater than 30% even if trees were not irrigated for up to 121 d. This may have been due to frequent precipitation and capillary rise to the root zone from the water table, located at 1 to 2 m below the soil surface, was sufficient to maintain adequate soil water even after several weeks of no irrigation.

92 For trees in containers in Krome very gravelly loam soil, SWD levels above 50% caused a reduction in ΨS that subsequently reduced gs. This correlated with a linear reduction in E and a sharp decline in A when gs fell below 50 mmol·m-2·s-1. Leaf gas exchange was better correlated with ΨS than with SWD level. Therefore, ΨS may be better indicator than SWD as determined from measurements of soil water content for irrigation scheduling of carambola trees in Krome soils.

93
SWD (%) 0 0 20 40 60 80 100

-1 ψS (MPa)

-2

-3

-4

Fig. 5-1. Soil water depletion (SWD) and stem water potential (Ψs) of carambola trees in containers in a greenhouse (y = -0.44 - (3.59e-06) x3, r2 = 0.82).

SWD (%) 0 0 20 40 60 80 100

-1
ψs (MPa)

-2

-3

-4

Fig. 5-2. Soil water depletion (SWD) and stem water potential (Ψs) of carambola trees in containers in the field (y = -0.48 - (2.79e-06) x3, r2 = 0.56).

94

200 180 160 140 gs (mmol m-2 s-1) 120 100 80 60 40 20 0 0 20 40 SWD (%) 60 80 100

Fig. 5-3. Soil water depletion (SWD) and stomatal conductance to water vapor (gs) of carambola trees in containers in a greenhouse (y = 95.46 – 0.0002 x3, r2 = 0.46).

95

200 180 160 140 120 100 80 60 40 20 -4 -3 -2 Ψs (MPa) -1 0 0 -2 -1 s )
-2 -1 gs (mmol m s )

Fig. 5-4. Stem water potential (Ψs) and stomatal conductance to water vapor (gs) of carambola trees in containers in a greenhouse (y = 11.88 + 132.94 ex , r2 = 0.54).

220 200 180 160 140 120 100 80 60 40 20 -4 -3 -2 ψs (MPa) -1 0 0

Fig. 5-5. Stem water potential (Ψs) and stomatal conductance to water vapor (gs) of carambola trees in containers in the field (y = 44.76 – 41.57 / x, r2 = 0.32).

gs (mmol m

96

4

2

1

-4

-3

-2 ψs (MPa)

-1

0

0

Fig. 5-6. Stem water potential (Ψs) and transpiration (E) of carambola trees in containers in a greenhouse (y = 0.30 + 2.94 ex, r2 = 0.55).

4

2

1

-4

-3

-2 ψs (MPa)

-1

0

0

Fig. 5-7. Stem water potential (Ψs) and transpiration (E) of carambola trees in containers in the field (y = 1.03 + 2.11 ex, r2 = 0.28).

-2 -1 E (mmol m s )

3

E (mmol m

-2 -1 s )

3

97

4

E (mmol m 2 s-1)

3

2

1

0

0

20

40

60

80

100

120

140

160

180

200

gs (mmol m 2 s-1)

Fig. 5-8. Stomatal conductance to water vapor (gs) and transpiration (E) of carambola trees in containers in a greenhouse (y = 0.33 + 0.02 x, r2 = 0.76).

4

E (mmol m 2 s-1)

3

2

1

0

0

20

40

60

80

100

120

140

160

gs (mmol m 2 s-1)

Fig. 5-9. Stomatal conductance to water vapor (gs) and transpiration (E) of carambola trees in containers in the field (y = 0.34 + 0.02 x, r2 = 0.75).

98

10 8 6 4 2 0 -2 A (ηmol m 2 s-1)

-4

-3

-2 ψs (MPa)

-1

0

Fig. 5-10. Stem water potential (Ψs) and net CO2 assimilation (A) of carambola trees in containers in a greenhouse (y = -0.09 + 9.76 ex, r2 = 0.70).

12 10 A (ηmol m 2 s-1) 8 6 4 2 0 -2

-4

-3

-2 ψs (MPa)

-1

0

Fig. 5-11. Stem water potential (Ψs) and net CO2 assimilation (A) of carambola trees in containers in the field (y = 1.19 - 2.95/x, r2 = 0.31).

99

10 8 A (ηmol m 2 s-1) 6 4 2 0 -2

0

20

40

60

80

100

120 -2 -1 s )

140

160

180

200

gs (mmol m

Fig. 5-12. Stomatal conductance (gs) and net CO2 assimilation (A) of carambola trees in containers in a greenhouse (y = -2.46 + 0.39 (lnx)2, r2 = 0.75).

10 8 A (ηmol m 2 s-1) 6 4 2 0 -2

0

20

40

60

80

100 -2 -1 s )

120

140

160

gs (mmol m

Fig. 5-13. Stomatal conductance (gs) and net CO2 assimilation (A) of carambola trees in containers in the field (y = -2.24 + 0.40 (lnx)2, r2 = 0.45).

100
ψL (MPa)
-4 -3 -2 -1 0 0

-1

-2

Field-container trees Greenhouse-container trees

-3

-4

Fig. 5-14. Leaf (ΨL) and stem water potentials (ΨS) of carambola trees in containers in the field (y = 0.10 + 1.00x, r2 = 0.97) and in a greenhouse (y = 0.33 + 0.91x, r2= 0.97).
ψS (MPa)
-0.9 -0.8 -0.7 -0.6 -0.5 -0.4 -0.3

-0.4

-0.6

-0.8

-1.0

-1.2

Fig. 5-15. Leaf (ΨL) and stem water potentials (ΨS) of carambola trees in an orchard in the field (y = 0.73 + 1.38x, r2 = 0.85).

ψL (MPa)

ψS (MPa)

CHAPTER 6 GROWTH, YIELD, AND FRUIT QUALITY OF CARAMBOLA AS AFFECTED BY SOIL WATER DEPLETION Carambola (Averrhoa carambola L.) is an evergreen, tropical fruit tree native to S.E. Asia where it is primarily cultivated. The tree is also cultivated in regions of Australia, India, Israel, the Caribbean Islands, and the United States (mainly Hawaii and Florida) (Crane, 1994; Mauro Pace, 2004, personal communication). A sweet type, ‘Arkin’, is the leading commercial carambola cultivar in Florida (Knight and Crane, 2002). In South Florida, carambola is grown in Krome very gravelly loam soil that has a high pH (7.4-8.4), low fertility, and low soil water holding capacity (Noble et al., 1996). Due to the hardness of the soil and to facilitate root growth, carambola trees are planted in perpendicular intersections of trenches that are approximately 50 to 75 cm wide and 50 cm deep (Colburn and Goldweber, 1961). In southern Florida, carambola growth ceases during winter and the tree remains quiescent until it refoliates in mid-March (NúñezElisea and Crane, 1998). Low temperatures, dry winds, and high light intensity (above 1000 µmol·m-2·s-1) cause carambola leaves to become chlorotic and trees partially defoliate from February to March (Núñez-Elisea and Crane, 1998; George et al., 2002a, 2002b). Fulfilling tree water requirements is important for economically profitable fruit production of carambola. Insufficient (Ismail and Noor, 1996a; Salakpetch et al., 1990) or excessive (Ismail and Noor, 1996b; Joyner and Schaffer, 1989) soil moisture can decrease tree growth and production. Increased water stress reduced vegetative growth

101

102 (Ismail et al., 1994, 1996) and flowering (Salakpetch et al., 1990) of carambola. Even when tree growth was not affected by drought, yield was significantly reduced in rainfed trees compared to irrigated trees in Malaysia, mainly due to a decrease in fruit size of non-irrigated trees (Bookeri, 1996). Little is known about water requirements and irrigation scheduling practices for carambola trees. In Australia, 30-75 mm of water per week was recommended for mature carambola trees grown in the Northern Territory (Lim, 1996) and northern Queensland (Galán Saúco et al., 1993). In southern Florida, Crane (1994) recommended 33 mm per ha twice a week during dry periods throughout the year. However, water application rates and frequencies that meet tree water requirements for optimum growth and yield have not been established. Understanding tree water requirements is important for proper irrigation management of carambola trees in subtropical climates such as that of South Florida. The objective of this study was to determine the effects of four levels of soil water depletion (SWD) on growth, yield and fruit quality of mature orchard-grown and young containergrown carambola trees in a Krome very gravelly loam soil. Materials and Methods Plant material and soil properties. The study was conducted in an orchard with 8-year-old ‘Arkin’ carambola trees grafted on open-pollinated Golden Star rootstock in Krome very gravelly loam soil in Homestead, Fla. at 25.5 oN Lat. and 80.5 oW Long. The trees were spaced at 4.5 m within rows and 6 m between rows and surrounded by an artificial (Polypropylene ribbon shade cloth) windbreak on the northern, eastern, and western perimeters and sapodilla [Manilkara zapota (L.) von Royen] trees as windbreaks on the southern perimeter of the orchard. Adjacent to the orchard, 2-year-old ‘Arkin’ carambola trees grafted on Golden Star rootstock were planted in 95 L plastic containers.

103 The containers were filled with Krome very gravelly loam soil which was excavated from a site next to the orchard. Fertilizer applications and pest management practices for the orchard and trees in containers followed recommendations for South Florida’s commercial carambola production (Crane, 1994). Soil water depletion treatments. Soil water depletion was determined by monitoring soil water content with multisensor capacitance probes (EnviroSCAN, Sentek PTY Ltd., Kent Town, Australia). Capacitance probes were installed 60 cm north of the trunk in the orchard and 20 cm from the trunk in containers of 3 trees per treatment. Each probe in the orchard consisted of four capacitance sensors located at soil depths of 10, 20, 30, and 50 cm. Probe sensors in containers were placed at of 10-, 20-, and 30-cm depths below the soil surface. The sensors recorded soil water content every 30 min and the data were stored in a datalogger and later downloaded to a portable computer for analysis. Description of the installation of the capacitance probe system in Krome soils was previously described by Al-Yahyai et al. (2003) and Núñez-Elisea et al. (2001), and technical specifications of the multisensor capacitance sensors were described by Paltineanu and Starr (1997). Data from the 10-, 20- and 30-cm depths sensors were summed and plotted using EnviroSCAN software (EnviroSCAN 4.0, Sentek PTY Ltd., Kent Town, Australia). Continuous measurements of soil water content using capacitance probes allowed for the determination of Field capacity (FC) (Hillel, 1998; Zekri and Parsons, 1999; Fares and Alva, 2000). Field capacity values were selected based on Veihmeyer and Hendrickson’s concept that define FC as the amount of water in the soil after excess water had drained and the rate of downward movement had decreased (Hillel 1998). After the treatment SWD levels were reached, irrigation was applied for each

104 treatment to restore soil moisture to FC in the orchard and to container capacity in containers using microsprinklers with a 360o wetting pattern (Maxijet, Dundee, Fla., USA) at 89 L·h-1. Soil water depletion levels were predetermined in the orchard and in containers based on a preliminary study (Al-Yahyai, unpublished data) whereby irrigation water was withheld and soil water content was monitored via capacitance probes until visual water stress symptoms (leaf yellowing and abscission) appeared on the trees. Irrigation of the orchard trees was initiated when SWD reached one of the following four levels (where 0% SWD = FC): 0-8% SWD, 9-11% SWD, 12-14% SWD, or 15-17% SWD. Irrigation of container trees was initiated when SWD reached one of the following four levels: 0-21% SWD, 22-31% SWD, 32-50% SWD, and 51-60% SWD. Containers were placed on metal bases above the ground. Soil water depleted faster in containers than in the orchard because lateral and capillary water movement into the container soils were prevented. Growth and yield measurements. Shoot growth was measured on four randomly selected, actively growing branches per tree in the orchard and three shoots per tree in the container study. The shoots were tagged at 10 cm below the shoot apex and measured using a metric ruler. Trunk caliper of trees in containers was measured from 27 July 2002 until 18 Dec. 2003 at 10 cm above the graft union. Fruit from carambola trees have an irregular shape and are composed of five to seven longitudinal ridges, thus fruit length was used to determine differences in fruit growth among treatments. Fruit length of four randomly selected fruit in the field and three fruit in containers were measured with a caliper. At the end of the container experiment, trees were removed from the containers

105 and the fresh and dry weights of the leaves, stems and roots of each tree were recorded to determine tree biomass. In the orchard, fruit from six trees per treatment were harvested during August (summer harvest) and December (winter harvest) of 2002 and 2003. The harvest dates coincided with commercial peak harvesting months for carambola trees in South Florida. Fruit from six trees per treatment were harvested on Dec. 2002 and Dec. 2003. Fruit were sorted based on the stage of maturity and color from 120 ohue (green) to 80 ohue (yellow) to immature (115 ohue), mature (108 ohue), and ripe (tree-ripened; 87 ohue) as determined with a colorimeter (Minolta Chroma Meter, Minolta, Inc., Ramsey, N.J.). Fruit were then counted and weighed to determine the total yield. Total soluble solids (TSS), fresh and dry weights of 20 randomly selected mature (108 ohue) and 20 ripe (87 ohue) fruit per tree were determined from six trees per treatment. Experimental design and statistical analysis. Treatments in the orchard and in containers were arranged in a completely randomized design. In the orchard, each treatment consisted of three rows with five trees per row. Two trees were randomly selected from each row for growth and yield measurements. In containers, treatments were replicated three times with two trees per replication. Data were analyzed by repeated measures analysis (SAS V.8.2, SAS Institute, Cary, NC) and treatment means were compared with Duncan’s multiple range test. Results and Discussion Shoot length. Shoot elongation from March to September in 2002 (Fig. 6-1A) and from April to December in 2003 (Fig. 6-1B) did not significantly differ among SWD treatments for carambola trees in the orchard. In orchard trees, this lack of shoot elongation in response to treatments could be attributed to sufficient soil water content

106 throughout the experiment. Frequent precipitation (1,355 mm in 2002 and 1,606 mm in 2003) and capillary rise from the water table located 1-2 m below soil surface may have provided an adequate water supply. In Malaysia, sufficient rainfall also resulted in no significant difference in shoot growth rate between irrigated and rain-fed field-grown carambola trees (Bookeri, 1996). Carambola trees can tolerate periods of low soil water content once they are established (Galán Saúco et al., 1993). Therefore, irrigation of carambola trees in an orchard at 17% SWD in Krome soils of South Florida did not reduce shoot length. Trunk diameter. Trunk diameter of 2-year-old carambola trees in containers in the 0-21% SWD treatment was higher than that of trees in the other treatments from July 2002 until Dec. 2003 (Fig. 6-2). Trees in the 0-21% SWD treatment had significantly greater trunk diameter than trees in the 32-50% SWD treatment throughout the experiment. Trunk diameter of trees in the 0-21% SWD treatment was significantly greater than that of trees in the 51-60% SWD treatments from September to December 2003. Trunk diameter of trees in the 22-31% SWD treatment was not significantly different from that of trees in the 32-50% SWD and 51-60% SWD treatments throughout the experiment. Trunk diameter of 2-year-old container-grown trees decreased when SWD was greater than 21%. Similarly, in a previous study, a smaller stem diameter was observed for water-stressed carambola trees in comparison to well-watered trees in containers (Ismail et al., 1994, 1996). Trunk diameter of container-grown carambola trees was smaller in treatments where SWD was allowed to drop below 32% compared to SWD of 0-21%, which indicated that whole-tree growth was reduced. Therefore, reduction in

107 trunk diameter was a better indicator of tree response to SWD than individual measurements of shoot length. Soil water depletion in the orchard, however, may not be high enough to cause a significant reduction in tree trunk diameter. Tree dry and fresh weight. There were no significant differences among treatments in leaf and root fresh and dry weights (Table 6-1) for container-grown trees at the end of the experiment. Trees in the 0-21% SWD treatment had significantly greater stem fresh weight than trees in the 22-31% SWD and 51-60% SWD treatments. Stem dry weight of trees in the 0-21% SWD treatment was also significantly greater than that of trees in the 51-60% SWD treatment. Total tree dry weight of trees irrigated at 0-21% SWD was significantly greater than for trees irrigated at the 51-60% SWD (Table 6-1). Stem and total dry weight reduction at high SWD levels are in agreement with studies from tropical regions such as of Malaysia in which leaf, stem, and root dry weights of carambola were reduced with increased water stress (Ismail et al., 1994,1996; Ismail and Noor, 1996a; Razi et al., 1992). There were no significant effect of SWD in the root:shoot ratio (on a dry weight basis) of trees in containers among SWD treatments (Table 6-1), due to the lack of SWD treatment effects on leaf and root dry weight. In contrast, Razi et al. (1992) found that root:shoot ratio of carambola was significantly higher in trees irrigated at 95% SWD in comparison to those irrigated at FC or 50% SWD. Ismail et al. (1996), however, reported a higher root:shoot ratio for carambola trees subjected to moderate water stress compared to well-watered trees or trees subjected to severe water stress. Severe water stress in the latter study may have decreased root growth due to reduced stomatal conductance and photosynthesis per unit leaf area (Hsiao, 1993; Chartzoulakis et al., 1993). An increased

108 root:shoot ratio in water-stressed trees is an adaptive mechanism to water stress resulting from greater allocation of dry matter to the roots than the shoots (Begg and Turner, 1976; Ismail et al., 1994; 1996; Razi et al., 1992). Young carambola trees grown in Krome soils outdoors in containers may not have reached water stress levels that could have resulted in a significant increase in the root: shoot ratio even in trees irrigated at 51-60% SWD. This was presumably due to the frequent precipitation during months when trees were actively growing (March to November). Fruit length and yield. Fruit length of carambola trees in the orchard did not differ significantly among SWD treatments (Fig. 6-3). Adequate soil water content during fruit growth periods in 2003 may have contributed to the lack of significant differences in fruit growth. Irrigation of carambola trees in Krome soil at 17% SWD did not reduce fruit length. There were no significant statistical interactions between SWD treatment and year or SWD treatment and harvest date for orchard or container-grown trees with respect to fruit number or weight. Number and weight of mature (108 ohue) and ripe (tree-ripened; 87 ohue) fruit from harvest 1 (Aug. 2002), harvest 2 (Dec. 2002), harvest 3 (Aug. 2003), and harvest 4 (Dec. 2003) of orchard trees were not significantly different among SWD treatments (Table 6-2). The total fruit number and weight were significantly greater in 2002 than in 2003. In addition, fruit number and weight from orchard trees in December were significantly greater than those harvested in August (data not shown). However, no significant differences in total fruit weight or number among treatments were observed in container-grown carambola trees.

109 In a study of carambola trees in Malaysia, Bookeri (1996) found that irrigation treatments did not affect fruit number but irrigated trees had fruit that were 17% larger than non-irrigated trees. In the present study, soil water content did not reach sufficiently low levels to cause water stress in orchard-grown trees and resulted in no significant differences in fruit length, number or weight among treatments. In 2-year-old containergrown carambola trees, the lack of yield differences among irrigation treatments may have been due to variability in fruit production among individual trees. Within the same treatment, several carambola trees in containers did not set fruit, while others produced 36 fruit per tree. Fruit fresh and dry weight of orchard-grown trees. There was a significant effect of year on fruit fresh and dry weights. Fruit from 2003 had higher fresh and dry weights than those from 2002. Fresh weight of mature fruit (108 ohue) of trees in the orchard was significantly lower in the 12-14% SWD treatment than that in 9-11% SWD treatment for fruit harvested in Aug. 2002 and significantly lower than 0-8% SWD treatment for fruit harvested in Dec. 2002 (Table 6-3). In Dec. 2003, fruit from trees in the 15-17% SWD had significantly greater fruit fresh weight than all other treatments, whereas fruit from trees in the 9-11% SWD had the lowest fresh weight. Dry weight of mature fruit from the Aug. 2002 harvest was significantly greater in the 9-11% and 15-17% SWD treatments than in the 0-8% and 12-14% SWD treatments (Table 6-4). In Dec. 2002, fruit of trees irrigated at 0-8% and 9-11% SWD had significantly greater fruit dry weight than those of trees in the 12-14% and 15-17% SWD treatments. In Aug. 2003, trees in 12-14% SWD had greater fruit dry weight than trees in

110 all other treatments. In Dec. 2003, fruit fresh weight was highest in the 0-8% SWD treatment and lowest in 12-14% SWD treatment. Fresh weight of ripe (tree-ripened, 87 ohue) fruit from the Aug. 2002 harvest was significantly greater in the 0-8% SWD treatment than in the 9-11% and 12-14% SWD treatments (Table 6-5). There were no significant differences in fruit fresh weight among treatments in Dec. 2002. In Aug. 2003, fresh weight of fruit from of trees in the 0-8% and 9-11% SWD treatments was significantly greater than that of the 12-14% SWD treatment. In Dec. 2003, fresh weight of ripe fruit from the 15-17% SWD treatment was significantly greater than that of trees in all other treatments, whereas, fruit from trees in the 9-11% SWD treatment had the lowest fruit fresh weight. In Aug. 2002, dry weight of ripe fruit from trees in the 15-17% SWD was significantly greater than that from all other treatments (Table 6-6). In Dec. 2002, fruit of trees in 15-17% SWD treatment had a lower fruit dry weight than fruit of trees in the 08% and 12-14% SWD treatments. In Aug. 2003, dry weight of ripe fruit was greatest in the 12-14% SWD treatment. Fruit dry weight in the 0-8% SWD was significantly greater than that of all other treatments. Fruit from 9-11% SWD and 12-14% SWD had the lowest dry weight in Dec. 2003. There was no effect of SWD on fruit fresh or dry weights of carambola trees in the orchard. Fruit fresh and dry weight of container-grown trees. Mature fruit had greater fresh weight than those in 2002 than in 2003. However, no significant statistical interaction between SWD treatment and year was observed for dry weight of ripe and mature fruit and for fresh and dry weights of ripe fruit. In Dec. 2002, mature fruit of trees in the 0-21% SWD treatment had a significantly greater fresh weight than those of trees

111 in all other treatments (Table 6-7). In Dec. 2003, fresh weight of mature fruit of trees in the 32-50% SWD treatment was significantly greater than that of trees in the 0-21% and 51-60% SWD treatments. Mature fruit dry weight was not significantly different among irrigation treatments in Dec. 2002. In Dec. 2002, dry weight of fruit from trees in the 2231% and 32-50% SWD treatments was significantly greater than that from trees in the 021% SWD treatment. Ripe fruit from container-grown trees in the 0-21% and 22-31% SWD treatments had greater fruit fresh weight than those in the 32-50% and 51-60% SWD treatments in Dec. 2002 (Table 6-8). In Dec. 2003, fresh weight of ripe fruit in the 22-31% and 32-50% SWD treatments was significantly greater than that in the 0-21% and 51-60% SWD treatments. In Dec. 2002, dry weight of ripe fruit was greater in the 32-50% and 51-60% SWD treatments than in the 0-21% and 22-31% SWD treatments (Table 6-8). In Dec. 2003, fruit dry weight was significantly greater in the 32-50% SWD than all other treatments. Total soluble solid (TSS) content. Total soluble solid (TSS) content of ripe (treeripened, 87 ohue) fruit from orchard trees varied among irrigation treatments from one harvest to another and significant interactions were found between treatments and year and treatments and harvest date for TSS. Total soluble solids of fruit harvested in 2003 were higher than those harvested in 2002 and TSS of fruit harvested in December was higher than in fruit harvested in August (data not shown). There was no clear trend of the effect of SWD treatments on fruit TSS. For example, fruit harvested in Dec. 2002 had the highest TSS in the 0-8% SWD treatment and lowest in the 15-17% SWD treatment, whereas, fruit harvested in Dec. 2003 had the highest TSS at the 15-17% SWD and

112 lowest TSS at 0-8% SWD (Table 6-9). Total soluble solid content in ripe fruit of trees in containers was not significantly different among treatments in Dec. 2002 (Table 6-10). However, fruit from Dec. 2003 had a significantly higher TSS in the 51-60% SWD treatment than in the 0-21% and 22-31% SWD treatments. Total soluble solid content was higher than the reported average TSS of 6.8-7.2% for carambola in southern Florida (Campbell and Koch, 1989; Crane et al., 1998). Increased TSS in fruit was perhaps due to reduced irrigation water applied in this study. Irrigation at 32-50% SWD and 51-60% SWD in containers resulted in an increase in TSS in Dec. 2003. Irrigation at high levels of SWD may be beneficial for increasing fruit TSS in containers. However, due to variability of TSS within harvest dates, no conclusion can be made on the effect of SWD treatments on ripe fruit TSS in orchard-grown carambola trees in this study. Conclusions Shoot growth of carambola trees grown in an orchard and in containers was not affected by SWD. In the orchard, sufficient soil water content from precipitation and the shallow water table possibly resulted in sufficient soil water content to obtain adequate vegetative growth and yields. For container-grown trees where lateral water movement and capillary rise were prevented, trunk diameter decreased in trees in the highest SWD treatment. Soil water depletion of 51-60% reduced total dry weight of trees in containers. The root:shoot ratio was also not significantly different among SWD treatments because there were no significant differences in leaf and root dry weights among SWD treatments. Reduction in dry weight of young carambola trees did not occur until soil water content decreased to 50% of FC. Fruit length, number and weight did not differ significantly among SWD treatments in orchard-grown trees. Total soluble solid content did not follow a specific trend in response to irrigation at various levels of SWD. Differences

113 among SWD treatments in mature and ripe fruit fresh and dry fruit weight were highly variable in container- and in orchard- grown trees. Thus it appears that irrigation of carambola trees in Krome very gravelly loam soil in an orchard can be applied at 17% SWD without a significant effect on fruit yield or quality.

114

Table 6-1. Effect of soil water depletion (SWD) on fresh and dry weights of leaves, stem, and roots of carambola trees in containers. Leaf fresh Leaf dry Stem fresh Stem dry Root fresh Root dry Total dry Root:shoot wt. (g) wt. (g) wt. (g) wt. (g) wt. (g) wt. (g) wt. (g)z ratioy 0-21% SWD 318.55 ax 98.89 a 2400.83 a 1032.79 a 2158.33 a 1005.29 a 2137.00 a 0.89 a 22-31% SWD 350.17 a 111.76 a 1824.00 b 820.29 ab 1947.00 a 909.29 a 1841.30 ab 0.97 a 32-50% SWD 433.20 a 153.22 a 1965.00 ab 873.29 ab 1767.00 a 805.29 a 1831.80 ab 0.82 a 51-60% SWD 229.36 a 67.89 a 1610.83 b 631.96 b 1552.50 a 728.62 a 1428.50 b 1.11 a z Total dry weight of leaves, stems and roots. y Root:shoot ratio based on dry weight. x Means within columns followed by the same letter are not significantly different according to Duncan’s multiple range test, P ≤ 0.05. Treatments 114

115 Table 6-2. Effect of soil water depletion (SWD) on total fruit number and fruit weight of carambola trees in an orchard. Treatments Totalz fruit no. Totalz fruit wt. (kg) 0-8% SWD 1107.3 ay 103.02 a 9-11% SWD 881.7 a 76.21 a 12-14% SWD 1123.3 a 112.33 a 15-17% SWD 1261.7 a 116.05 a z Total of four harvests per tree (Aug. 2002, Dec. 2002, Aug. 2003, and Dec. 2003) of fruit excluding immature fruit. y Means within columns followed by the same letter are not significantly different according to Duncan’s multiple range test, P ≤ 0.05.

116 Table 6-3. Effect of soil water depletion (SWD) on fresh weight of mature fruit (108 o hue) of carambola trees in an orchard. Fruit fresh wt. (g) Dec. 2002 Aug. 2003

121.63 ab 127.21 a 125.59 a 147.42 b 9-11% SWD 129.83 a 120.40 ab 122.73 a 81.02 c 12-14% SWD 117.60 b 111.48 b 116.78 a 147.79 b 15-17% SWD 123.74 ab 119.90 ab 115.54 a 162.72 a z Means within columns followed by the same letter are not significantly different according to Duncan’s multiple range test, P ≤ 0.05.

Treatments 0-8% SWD

Aug. 2002
z

Dec. 2003

117 Table 6-4. Effect of soil water depletion (SWD) on dry weight of mature fruit (108 ohue) of carambola trees in an orchard. Fruit dry wt. (g) Treatments Aug. 2002 Dec. 2002 Aug. 2003 Dec. 2003 z 0-8% SWD 11.89 b 11.23 a 17.86 b 15.21 a 9-11% SWD 15.45 a 10.56 a 16.64 b 13.20 bc 12-14% SWD 12.34 b 9.11 b 20.22 a 12.31 c 15-17% SWD 15.56 a 9.49 b 17.64 b 14.33 ab z Means within columns followed by the same letter are not significantly different according to Duncan’s multiple range test, P ≤ 0.05.

118 Table 6-5. Effect of soil water depletion (SWD) on fresh weight of ripe fruit (treeripened, 87 ohue) of carambola trees in an orchard. Fruit fresh weight (g) Dec. 2002 Aug. 2003

163.17 a 129.44 a 154.46 a 182.84 b 9-11% SWD 141.55 b 124.78 a 155.64 a 96.61 c 12-14% SWD 143.95 b 131.09 a 137.96 b 180.19 b 15-17% SWD 151.00 ab 124.07 a 146.13 ab 202.85 a z Means within columns followed by the same letter are not significantly different according to Duncan’s multiple range test, P ≤ 0.05.

Treatments 0-8% SWD

Aug. 2002
z

Dec. 2003

119 Table 6-6. Effect of soil water depletion (SWD) on dry weight of ripe fruit (tree-ripened, 87 ohue) of carambola trees in an orchard. Fruit dry weight (g) Aug. 2002 Dec. 2002 Aug. 2003 Dec. 2003 Treatments z 0-8% SWD 14.70 b 11.97 a 21.00 b 22.85 a 9-11% SWD 14.02 b 10.93 ab 21.30 b 17.24 c 12-14% SWD 14.65 b 11.41 a 23.77 a 16.37 c 15-17% SWD 17.02 a 10.17 b 21.33 b 19.75 b z Means within columns followed by the same letter are not significantly different according to Duncan’s multiple range test, P ≤ 0.05.

120 Table 6-7. Effect of soil water depletion (SWD) on fresh and dry weights of mature fruit (108 ohue) of carambola trees in containers. Fruit fresh wt.(g) Fruit dry wt.(g) Treatments Dec. 2002 Dec. 2003 Dec. 2002 Dec. 2003 z 52.58 b 9.39 a 4.20 b 0-21% SWD 105.61 a 22-31% SWD 87.40 b 67.95 ab 9.81 a 7.67 a 32-50% SWD 84.19 b 87.50 a 9.67 a 7.51 a 51-60% SWD 80.84 b 53.85 b 9.64 a 5.60 ab z Means within columns followed by the same letter are not significantly different according to Duncan’s multiple range test, P ≤ 0.05.

121 Table 6-8. Effect of soil water depletion (SWD) on fresh and dry weights of ripe fruit (tree-ripened, 87 ohue) of carambola trees in containers. Fruit fresh wt.(g) Fruit dry wt.(g) Treatments Dec. 2002 Dec. 2003 Dec. 2002 Dec. 2003 z 0-21% SWD 108.69 a 51.55 b 9.62 b 4.46 b 22-31% SWD 113.56 a 93.72 a 9.89 b 5.65 b 32-50% SWD 53.34 b 97.68 a 11.55 a 9.47 a 51-60% SWD 56.05 b 62.25 b 11.23 a 5.98 b z Means within columns followed by the same letter are not significantly different according to Duncan’s multiple range test, P ≤ 0.05.

122

Table 6-9. Effect of soil water depletion (SWD) on total soluble solid (TSS) content of ripe fruit (tree-ripened, 87 ohue) of carambola trees in an orchard. TSS (%) Treatments Aug. 2002 Dec. 2002 Aug. 2003 Dec. 2003 z 0-8% SWD 9.31 a 10.43 c 7.45 c 9.13 a 9-11% SWD 9.33 a 10.68 bc 8.36 b 8.73 b 12-14% SWD 9.04 a 11.04 b 9.00 a 8.34 c 15-17% SWD 8.99 a 11.97 a 8.08 b 8.13 d z Means within columns followed by the same letter are not significantly different according to Duncan’s multiple range test, P ≤ 0.05.

123

Table 6-10. Effect of soil water depletion (SWD) on total soluble solid (TSS) content of ripe fruit (tree-ripened, 87 ohue) of carambola trees in containers. TSS (%) Treatments Dec. 2002 Dec. 2003 z 0-21% SWD 8.11 a 10.61 c 22-31% SWD 8.14 a 10.83 bc 32-50% SWD 7.86 a 11.28 ab 51-60% SWD 7.45 a 11.44 a z Means within columns followed by the same letter are not significantly different according to Duncan’s multiple range test, P ≤ 0.05.

124

50

A
40

Shoot length (cm)

30

20 0-8% SWD 9-11% SWD 12-14% SWD 15-17% SWD

10

0

B
50

Shoot length (cm)

40

30

20

10

0

01 -M ay

01 -A ug

Date

Fig. 6-1. Effect of soil water depletion (SWD) on shoot length of orchard-grown carambola trees during 2002 (A) and 2003 (B). Symbols with vertical bars represent means ± SE of 4 shoots per tree for 6 trees.

01 -N ov

01 -S e

01 -D ec

ar

01 -A pr

01 -J un

01 -M

01 -O ct

01 -J ul

p

125

50 0-21% SWD 22-31% SWD 32-50% SWD 51-60% SWD

45

Trunk diameter (mm)

40

35

30

25

20

Date Fig. 6-2. Effect of soil water depletion (SWD) on trunk diameter of young carambola trees grown in containers. Symbols with vertical bars represent means ± SE of 6 trees.

-0 2 01 -S ep -0 2 01 -N ov -0 2 01 -J an -0 3 01 -M ar -0 3 01 -M ay -0 3 01 -J ul -0 3 01 -S ep -0 3 01 -N ov -0 3

01 -J ul

126

100 90 0-8% SWD 9-11% SWD 12-14% SWD 15-17% SWD

Fruit length (mm)

80 70 60 50

Summer harvest
40 30

Winter harvest

03

01 -J un -0 3

01 -O ct -0 3

-J

ug -

ep -0 3

ov -0 3

Date Fig. 6-3. Effect of soil water depletion (SWD) on fruit length of orchard-grown carambola trees during 2003 summer harvest (August) and winter harvest (December). Symbols with vertical bars represent means ± SE of 4 fruit per tree for 6 trees.

01 -N

01 -A

01 -S

01 -D

01

ec -0 3

ul -0 3

CHAPTER 7 PHENOLOGICAL CYCLES OF CARAMBOLA AT FOUR LEVELS OF SOIL WATER DEPLETION IN SOUTH FLORIDA The phenological cycle describes the morphological developmental pattern of tree crops (Cull, 1986). In perennial evergreen trees such as carambola, the annual phenological cycle consists of multiple factors operating in synchrony, producing multiorder interactions that influence productivity (Cull, 1986). This pattern depends on species, cultivar, management, and environment. Environmental factors, however, are considered the most important factors that affect tree performance and management strategies (Whiley et al., 1988). The phenological cycle of tropical evergreen trees can be divided into three phases: the vegetative phase, the flower development phase, and the root development phase (Cull, 1986). Each phase has an effect on the other phases. For example, vegetative growth is essential for carbohydrate accumulation that is required for the development of young, photosynthetically efficient leaves, root growth, vegetative flushes, and fruit growth and development (Cull, 1986). The phases of the phenology cycle of tropical fruit trees are influenced by climatic conditions, mainly temperature and precipitation (Cull, 1986). The effects of climatic and environmental conditions on annual phenological cycles have been studied in several tropical and subtropical fruit crops including avocado (Cull, 1986; Salomon, 1984; Whiley et al., 1988), citrus (Salomon, 1984), mango (Whiley, 1993), and olive (Cimato et al., 1990). Just as the phenology of deciduous temperate trees is mainly determined by seasonal variations in temperature and

127

128 photoperiod, the phenology of tropical fruit is affected by seasonal variability in water availability (Reich and Borchert, 1984). Phenological cycles of tropical evergreen trees are less obvious than those of temperate fruit trees. Tropical fruit growers must recognize annual critical growth changes in order to utilize growth cycles as orchard management tools (Whiley et al., 1988). Environment and orchard cultural practices affect vegetative and reproductive development throughout the annual phenological cycle of carambola. The effect of several environmental factors on carambola have been studied including low temperatures (Campbell and Malo, 1981; Campbell et al., 1985; Crane, 1994; George et al., 2002a, 2002b; Watson et al., 1988), drought (Ismail and Awang, 1992; Ismail and Noor, 1996a; Ismail et al., 1994, 1996; Salakpetch et al., 1990) , flooding (Ismail and Noor, 1996b; Joyner and Schaffer, 1989), sunlight, (Marler et al., 1994), and wind (Campbell, 1985; Crane, 1993; Crane et al., 1994a; Marler and Zozor, 1992). In addition to environmental factors, cultural practices such as cultivar selection (Salakpetch et al., 1990), mulching (George et al., 2000; 2001), fertilization (Campbell, 1985,1989; Crane, 1994; Knight, 1982), and pruning (Crane et al., 1991; Núñez-Elisea and Crane, 1998, 2000) also influence vegetative and reproductive growth of carambola trees. In their native, tropical habitat, carambola trees grow, flower, and fruit year-round (Núñez-Elisea and Crane, 2000). Trees may produce new flushes, flowers, and immature and mature fruit simultaneously (Galán Saúco et al., 1993). Under subtropical conditions such as those in south Florida, canopy development is poor and trees defoliate from February to March presumably due to low temperatures, dry winds, and high light intensity (Núñez-Elisea and Crane, 1998). Trees begin to refoliate from March to midMay and may continue to produce new leaves until mid-October (Núñez-Elisea and

129 Crane, 1998). Shoots of carambola alternate growth with periods of quiescence. Apical growth results in an extension of the shoot axis, with each growth flush containing between eight and 12 leaves (Núñez-Elisea and Crane, 1998). Shoot flushing is enhanced by warm weather and rain. Shoots formed during the spring flush about four to five times per growth season but those formed in the late summer normally flush once before shoot extension growth ceases in the fall (Núñez-Elisea and Crane, 1998). Carambola has several flowering periods and may flower continuously during the year under favorable environmental conditions (Núñez-Elisea and Crane, 1998, 2000). The carambola production season in south Florida extends from mid-July to midFebruary (Campbell et al., 1985; Núñez-Elisea and Crane, 1999, 2000). Two reproductive growth phases occur in south Florida. The first phase is characterized by a flowering peak in May and with fruit maturing from July to October. The second phase has a peak bloom in September with fruit maturing from December to February (Campbell et al., 1985; Núñez-Elisea and Crane, 1998). These flowering phases and the ability of different wood types to flower indicate that floral initiation is affected by environmental, physiological, and cultural factors (Núñez-Elisea and Crane, 2000). Manipulation of growth and production cycles, therefore, requires an understanding of developmental and phenological properties of the tree. Soil water content is among the three major growth and productivity determinants (in addition to temperature and nutrition) that can be regulated throughout the phenological cycles of tropical fruit crops (Cull, 1986). A few studies have investigated the effect of soil water depletion (SWD) on the phenology of tropical fruit crops. For example, water requirements of avocado trees in relation to the growth cycle were studied

130 by Whiley et al. (1988). Núñez-Elisea and Crane (1998) studied the phenological cycles of carambola trees in South Florida. However, little is known about the response of carambola phenological cycles to SWD in Krome very gravelly loam soil in a subtropical climate like Florida. This study was carried out to determine the effect of four levels of SWD on the phenological cycle of carambola trees grown in the subtropical climate of South Florida. Materials and Methods Experimental site and plant material. In an orchard, 8-year-old ‘Arkin’ carambola trees grafted on open-pollinated Golden Star rootstock were visually monitored for phenological growth cycles throughout 2003. The trees were planted at 4.5 m within and 6.0 m between rows and were covered by an artificial (polypropylene ribbon shade cloth) windbreak on the northern, eastern, and western perimeters and sapodilla [Manilkara zapota (L.) von Royen] trees on the southern perimeter of the orchard. All the trees were topped to a 2.7 m height on 23 Apr. 2003. The orchard was located at the Tropical Research and Education Center in Homestead, Fla. at 25.5 oN Lat. and 80.5 oW Long. Average monthly temperatures in 2003 were lowest in January (17.9 oC) and the highest (30.4 oC) in July (Fig. 7-1), which are near the ideal temperature range for growth of carambola (21 to 32 oC) (Ngah et al., 1989). Historical climatic records of Homestead, Fla. for 41 years showed an average annual precipitation of 1,571 mm, 79% of which occurred from May through October (Fig. 7-1). The same pattern occurred during 2003, when 74% of the 1,606 mm total precipitation occurred during this period. The soil at the experimental site is Krome very gravelly loam [loamy-skeletal, carbonatic, hyperthermic, Lithic Udorthents (Noble et al.,

131 1996)]. Trees were planted at the intersection of perpendicular trenches that were approximately 50 to 75 cm wide and 50 cm deep (Colburn and Goldweber, 1961). A second experiment was also conducted during the same period in 2003 to study the phenological cycle of 2-year-old ‘Arkin’ carambola trees grafted on Golden Star rootstock planted in 95 L containers on 4 June 2002. The containers were filled with Krome very gravelly loam soil and placed in the field adjacent to the orchard used for the phenological study of mature trees. Fertilizer and pest management were in accordance with a standard practice for commercial carambola production in south Florida (Crane, 1994). Soil water depletion treatments. Soil water depletion was determined by monitoring soil water content with multisensor capacitance probes (EnviroSCAN, Sentek PTY Ltd., Kent Town, Australia). Three probes per treatment were installed 60 cm north of the trunk of trees in the orchard and 20 cm from the trunk of trees in containers. Each probe in the orchard contained four capacitance sensors located at 10, 20, 30, and 50 cm depths in the soil. Sensors were placed at 10, 20, and 30 cm depths in containers. The sensors recorded soil water content every 30 min. The data were stored in a datalogger and later downloaded to a portable computer for analysis. Installation of the system in Krome soils was described previously by Al-Yahyai et al. (2003) and Núñez-Elisea et al. (2001), and technical specifications of the multisensor capacitance probes were discussed by Paltineanu and Starr (1997). Data from the 10, 20 and 30 cm depth sensors were averaged and plotted using EnviroSCAN software (EnviroSCAN 4.0, Sentek PTY Ltd., Kent Town, Australia).

132 Field capacity (FC) values were determined based on the amount of water in the soil after excess water had drained and the rate of downward movement had decreased. Trees in the orchard and in containers were irrigated with microsprinklers (Maxijet, Dundee, Fla., USA) which had a discharge rate 89 L·h-1 and a 360o wetting pattern. Levels of soil water depletion were predetermined in the orchard and in containers based on a preliminary study (Al-Yahyai, unpublished data) whereby irrigation water was withheld and soil water content was monitored with capacitance probes until visual water stress symptoms (leaf yellowing and abscission) appeared on the trees. Containers were placed on metal bases above the ground to prevent capillary movement of water from the soil to the containers. Irrigation of the orchard trees was initiated when SWD reached one of the following four levels (where 0% SWD = FC): 0-8% SWD, 9-11% SWD, 12-14% SWD, or 15-17% SWD. Irrigation of container trees was initiated when SWD reached one of the following four levels: 0-21% SWD, 22-31% SWD, 32-50% SWD, and 51-60% SWD. Phenological observations. Orchard and container-grown trees were visually evaluated weekly for 37 weeks in 2003. Observations of vegetative and reproductive development included: 1) shoot flushing: shoot growth following bud break or resumption of growth of quiescent shoots. Following the initial post-winter shoot flush, observations were made on new axillary bud break (George et al., 2000); 2) shoot growth: the relative number of continuously growing axillary shoots; 3) flowering: growth of the panicle from fully developed buds to flower opening (bloom) (Whiley et al., 1988). Most panicles were formed on branches on the periphery of the canopy, and quiescent buds on older branches. Flowers were also formed on scaffold branches and on

133 the main trunk (Núñez-Elisea and Crane, 2000); 4) fruit load: the relative number of fruit per tree. Fruit from six trees per treatment were completely harvested on 18-22 Aug. (summer harvest) and on 15-19 Dec. (winter harvest), which coincided with the peak harvesting season in south Florida for 2003. Experimental design and statistical analysis. Treatments in the orchard and containers were arranged in a completely randomized design. Each treatment consisted of three rows. In the orchard, the numbers of trees per treatment were 15 for 0-8% SWD, 11 for 9-11% SWD, 11 for 12-14% SWD, and 14 for the 15-17% SWD treatment. For containers, each treatment was replicated three times with two trees per replication. The lack of uniformity in replications among treatments in the orchard was due to some rows containing unhealthy trees or support posts for the artificial windbreak that resulted in an uneven number of trees per row. Trees were evaluated based on a score of 0 to 100. Ranking were based on the following rating: 0 (none), 1 (very low), 2 (low), 3 (medium), 4 (high), and 5 (very high to maximum). A similar approach was previously used to evaluate carambola canopy leaf color (George et al., 2000) and flowering (Crane et al., 1991; George et al., 2001). The ranked data were analyzed using the Kruskal-Wallis nonparametric test for unequal population sample size (Miller, 1980; Neter et al., 1990). Data were analyzed using the NPAR1WAY procedure in SAS (SAS Institute, Cary, N.C.). Multiple mean comparisons were performed as described by Miller (1980) and Neter et al. (1990). Results Shoot flush. In trees in the orchard, shoot flushing began during the last week of February and reached its peak by mid-March (Fig. 7-2A.). After the initial shoot flush in the spring, trees continued to flush but at a low rating until the third week of October.

134 Another small increase in shoot flushing occurred following harvest at the end of September and peaked at the beginning of October. The highest average spring flush rating, 4.6, occurred in the middle of March for trees in the 9-11% SWD treatment. The fall flushing was highest for trees in the 12-14% SWD treatment with an average rating of 1.8 in the first week of October. There were no significant differences in shoot flushing among treatments during the peak spring and fall shoot flushes. Shoot flushing during May and June occurred after trees were topped on 23 Apr. Shoot growth ceased toward the end of October and trees became quiescent during November and December. There was a significant difference in shoot flushing on 28 Feb. between trees in the 0-8% SWD that had a high rating (3.8) and those in the 12-14% SWD treatment with a low rating (2.8). There were no significant differences in shoot flushing ratings among the other treatments (Table 7-1). There were also no significant differences in shoot flushing ratings among treatments for the rest of the year. Shoot growth of container-grown carambola trees began during the third week of February (Fig. 7-2B). However, unlike orchard trees, shoot flushing of container-grown trees fluctuated throughout the year and no bud break was observed from mid-May to mid-July and from mid-November until the end of December. Trees in the 0-21% SWD and 22-31% SWD treatments had higher shoot flushing ratings than trees in the 32-50% and 51-60% SWD treatments from mid-February to mid-May. Container-grown trees started a new shoot flush in August that reached a peak by the first week of September. Shoot flushing continued at a low level until it ceased by the second week of November. Significant differences were observed on 25 Apr. between trees in the 0-21% SWD treatment that had a low shoot flushing rating of 1.7 and trees in the 22-31% SWD

135 treatment that had no shoot flushes (Table 7-2). Although shoot flushing varied among treatments, no further significant differences among irrigation treatments were observed. Extension shoot growth. Shoot extension of carambola trees in the orchard was observed from the first week of March until the third week of December. Extension shoot growth was very rapid at the beginning of the growing season in March with a gradual decrease in growth by the end of September followed by a rapid decline from October to November until growth ceased toward the end of December (Fig. 7-3A). Shoot extension ratings of the axillary shoots fluctuated from March to September. During the initial growth of the axillary shoots of trees in the orchard in March and April, there were no significant differences among treatments (Table 7-1). However, trees in the 9-11% SWD treatment had a significantly higher shoot growth rating than trees in the 12-14% SWD treatment on 23 May, 30 May, 7 June, and 13 June. Also significantly more shoot growth occurred in trees in 0-8% SWD than in the 12-14% SWD treatment. In addition, trees in the 0-8% and 9-11% SWD treatments had significantly higher shoot growth rating than trees in the 12-14% SWD treatment on 8 Aug. No significant differences in shoot growth rating were observed between trees irrigated at 0-8% and 1517% SWD (Table 7-1). Container-grown trees had similar shoot growth patterns to those of orchard trees throughout the year (Fig. 7-3B). Shoot growth of container-grown trees was high during the last week of March and the first week of April. Unlike orchard trees, however, shoot growth ratings of trees in containers did not gradually decrease until September (Fig. 73B). Trees in the 32-50% and 51-60% SWD treatments generally had fewer shoots growing than trees in the 0-21% or 22-31% SWD treatments.

136 Carambola trees in the 0-21% SWD had significantly fewer axillary shoots growing than trees in the 51-60% SWD treatment on 25 Apr. (Table 7-2). However, trees in the 0-21% SWD treatment had more axillary shoots growing than trees in the 32-50% SWD treatment on 30 May and 7 June, and the 51-60% SWD treatment on 19 Sept. Axillary shoot growth ratings of these trees were significantly higher than for trees in the 32-50% SWD treatment (Table 7-2) on 30 May and 7 June. Shoot growth ratings of trees in the 0-21% SWD treatment were significantly higher than that of trees in the 51-60% SWD treatment. Flowering. Flowering of trees in the orchard was first observed in the first week of March (Fig. 7-4A). However, fruit set from this bloom was very low. During the second half of April, flowering increased and reached its peak during the last 2 weeks of May. Flowering then continued at a moderate level until the fruit were harvested during the third week of August. Following harvest, flowering peaked again from mid-September to mid-October. The amount of flowers decreased following this period and flowering ceased by the end of December (Fig. 7-4A). Trees in the 9-11% SWD treatment had a significantly higher flowering rating than trees in the 15-17% SWD treatment on 1 Oct. (Table 7-1). On 17 Oct., trees in the 9-11% SWD treatment had the highest flowering rating (Table 7-1). The first flowering peak in container trees occurred in March and flowering declined toward the end of April (Fig. 7-4B). Low flowering ratings were observed from mid-May to the end of July. An increase in flowering began in August which reached its peak by the end of October, and then declined by December (Fig. 7-4B). Containergrown trees had lower flowering ratings during the summer and fall compared to that of

137 orchard trees. On 7 and 14 Mar., trees in the 0-21% SWD treatment had a significantly higher flowering rating than trees in the 51-60% SWD treatment (Table 7-2). Fruiting. In the orchard, fruit that were not harvested in the winter remained on the tree until March when they ripened and abscised (Fig. 7-5A). Trees did not set fruit from newly formed flowers until the second half of April, with a sharp increase in fruit set during the first two weeks of May. Fruit production continued as new flowers produced and set fruit for the rest of the year. The major fruit harvest periods were August and December. However, trees continuously produced fruit during the year. Orchard trees in the 15-17% SWD treatment had a significantly higher fruit number rating on 30 May than trees in the 12-14% SWD treatment but fruit number ratings in the 15-17% SWD and 12-14% SWD treatments did not differ from that in the 9-11% SWD and 0-8% SWD treatments (Table 7-1). Fruiting was erratic and there was no harvestable yield for carambola trees in containers until October with a peak yield in mid-November (Fig. 7-5B). During the first flowering phase there were no harvestable fruit in August, resulting in only one harvest in December. Fruit number ratings were not significantly different among treatments throughout the year. This was attributed to the low number of fruit produced and the variation in fruit yield among container-grown trees in 2003. Discussion In the subtropical climate of South Florida, carambola trees did not produce shoot flushes until mid-February in 2003. Adverse climatic conditions including low temperatures and dry winds were cited as the reason for this lack of shoot development during that time of year in south Florida (Campbell et al., 1985; George et al., 2002a, 2002b; Núñez-Elisea and Crane, 1998, 2000) in comparison to tropical climates where

138 carambola continuously produce new shoots (Núñez-Elisea and Crane, 2000; Galán Saúco et al., 1993). The peak amount of shoot flushing occurred in the spring following a period of quiescence and a second peak of shoot growth flush occurred in the fall following the summer harvest, before shoot growth ceased in the winter. Continuous shoot growth flushing throughout the spring, summer, and winter was due to an alteration of growth and quiescence as observed by Núñez-Elisea and Crane (1998). Following topping of orchard trees in April, pruned branches produced 70% of the new growth in May and June, whereas container-grown trees did not flush during this period. Pruning of carambola increased vegetative shoot flushing (Núñez-Elisea and Crane, 2000). Shoot flushing of carambola trees in the orchard and in containers was not affected by irrigation at four levels of SWD under the subtropical climatic conditions of south Florida in 2003. The lack of response was presumably due to sufficient soil water from precipitation (Fig. 7-1) in the orchard and in containers. In addition, capillary rise of water from the high water table (1-2 m below soil surface) to the root zone may have minimized SWD treatment effects on orchard trees. Overall shoot growth was not affected by SWD treatments in the orchard. This lack of response was due to soil water depletion not reaching sufficiently low levels to cause a reduction in shoot growth. In Malaysia, Bookeri (1996) also found no response of overall growth of carambola trees in an orchard to irrigation due to sufficient rainfall following a drought. Similarly, Núñez-Elisea and Crane (1998) reported that, in southern Florida, vegetative growth of carambola occurred throughout the spring and summer months in the presence of warm temperatures and rain. For container trees where lateral and capillary soil water movement into the root zone was restricted, trees irrigated at field

139 capacity had a significantly higher extension shoot growth than trees in the 32-50% and 51-60% SWD treatments. A reduction of shoot growth of fruit crops in response to water stress is well documented (Hsiao and Acevedo, 1974; Lakso, 1985). A reduction in vegetative growth in response to water stress was reported previously for containergrown (Ismail et al., 1994; Ismail and Noor, 1996a) and field-grown carambola trees (Ismail et al., 1996). Flowering of carambola trees in an orchard and in containers in South Florida in 2003 occurred in March. Flowering in March may have resulted from above average temperatures in February and March. The average monthly temperature in February was 22.3 oC (annual average is 19.9 oC) and maximum temperature was 30.8 oC. In March, the average monthly temperature was 25.9 oC (annual average is 22.1 oC) and maximum temperature was 32.3 oC. In the subtropical climate of south Florida, George et al. (2001) found that flowering of carambola trees in an orchard increased in response to increased soil temperature. In contrast, flowering of young container-grown carambola was less affected by temperature than by water stress in a greenhouse experiment in Australia (Salakpetch et al., 1990). For orchard trees in our study, two flowering periods were observed, the first was from April to June with a peak in May, and the second flowering period was from August to October with a peak in September. These two flowering peak periods in South Florida were also observed by Campbell et al. (1985) and Núñez-Elisea and Crane (1998). Under optimum environmental conditions, however, carambola has several flowering periods throughout the year (Samson, 1980; Tidbury, 1976; NúñezElisea and Crane, 1998; 2000).

140 Orchard trees were able to sustain more flowers during the first flowering period (April to June) than container trees. It can be speculated that due to the limited carbohydrate reserves in young container trees following initial flowering in March, flowering ratings were very low in the first flowering period from April to June. At critical stages of the growth cycle, competition for carbohydrate reserves between reproductive and vegetative growth leads to depletion of these reserves (Whiley et al., 1988). In container trees, vegetative growth may have taken precedence over reproductive growth to replenish these carbohydrate reserves (Anuar et al., 1992), which explains the absence of the first flowering phase in container-grown carambola trees. Overall carambola flowering in orchard and container-grown trees was not affected by SWD levels except for two observation dates when trees receiving the least amount of water had the fewest flowers. The inconsistent response of flowering of carambola trees to SWD in this study was similar to observations for avocado, lychee and mangoes. In avocado and lychee, Chaikiattiyos et al. (1994), reported that water stress reduced vegetative growth but did not induce flowering. Flowering of mango in response to water deficit was also found to be inconsistent, where trees in the orchard flowered but those in a greenhouse did not flower following rewatering (Whiley, 1993). Fruit production of carambola trees was not influenced by soil water depletion levels. This lack of response is contrary to what was found in lychee (Menzel et al., 1995), where fruit set and growth was reduced by water stress. Adequate soil water for fruit growth and development of carambola trees in an orchard in south Florida resulted in no differences in fruit number among SWD treatments. Fruit production of container-

141 grown trees was negligible and varied greatly, thus no significant differences were observed among treatments. Conclusions Phenological cycles of carambola trees including shoot flushing, extension shoot growth, flowering, and fruiting showed little response to irrigation at four SWD levels. This lack of response was likely caused by sufficient soil water due to precipitation and capillary rise from the high water table located about 1-2 m below the soil surface. Environmental factors such as temperature and cultural practices such as pruning appeared to have altered the annual phenological cycle by influencing vegetative and reproductive growth. The phenological cycles of young trees grown in containers were not as well-defined as those of mature trees in an orchard. The response of containergrown trees to irrigation varied among trees rather than among treatments. Further studies over longer periods are needed to better understand the phenology of these trees. Nonetheless, information on phenology of carambola in this study may provide useful guidelines for orchard management in South Florida.

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Table 7-1. The effect of soil water depletion (SWD) on shoot flushing, axillary shoot growth, flowering, and fruit load of orchardgrown carambola trees in 2003. Ratings Shoot flushing Treatments 0-8% SWD 9-11% SWD 12-14% SWD 15-17% SWD
z y

Axillary shoot growth 23-May 30-May 4.6 ab 4.7 a 3.9 b 4.5 ab 4.5 ab 4.8 a 4.1 b 4.5 ab 7-Jun 4.4 ab 4.8 a 3.9 b 4.1 ab 13-Jun 4.3 ab 4.7 a 3.6 b 3.7 b 3-Jul 4.6 a 4.4 ab 3.7 b 3.9 ab 8 Aug. 3.9 a 3.8 a 3.2 b 3.3 ab

Flowering 1 Oct. 17 Oct. 4.1 ab 4.6 a 3.8 ab 3.5 b 3.2 b 3.8 a 3.0 b 3.0 b

Fruit load 30-May 2.9 ab 3.3 ab 142 2.7 b 3.6 a

28 Feb.z 3.8 ay 3.7 ab 2.8 b 3.4 ab

Dates listed when significant differences among treatments were detected (total number of observation dates was 37). Different letters within columns indicate significant differences among means according to the Kruskal-Wallis non-parametric test, α ≤ 0.1.

143

Table 7-2. The effect of soil water depletion (SWD) on shoot flushing, axillary shoot growth, and flowering of container-grown carambola trees in 2003. Ratings Shoot flushing Treatments 0-21% SWD 22-31% SWD 32-50% SWD
z

Axillary shoot growth 25 Apr. 30-May 3.3 b 4.8 ab 4.6 ab 4.8 a 4.4 ab 3.4 b 7-Jun 4.8 a 4.4 ab 3.2 b 19 Sept 8 Aug. 4.0 a 3.8 ab 3.6 ab 2.8 a 2.0 ab 1.4 ab

Flowering 7 Mar. 14 Mar. 2.8 a 2.0 ab 1.4 ab 3.2 a 2.2 ab 1.8 ab 143

25 Apr.z 1.7 ay 0.0 b 1.4 ab

51-60% SWD 0.3 ab 5.0 a 4.2 ab 4.0 ab 3.0 b 1.2 b 1.2 b 1.0 b Dates listed if significant differences among treatments were detected (total number of observation dates was 37). y Different letters within columns indicate significant differences among means according to the Kruskal-Wallis non-parametric test, α ≤ 0.1.

144

500 450 400 2003 average monthly rainfall 41 yr average monthly rainfall 2003 average monthly temp.

35 30 25 20 15 10 5 0 Jan Feb Mar April May June July Aug Sept Oct Nov Dec Temperature (oC)

Precipitation (mm)

350 300 250 200 150 100 50 0

Month

Fig. 7-1. Average 41-year and 2003 monthly precipitation and average 2003 monthly temperature in Homestead, Fla. Source: Florida Automatic Weather Network (FAWN) (2003) and The National Oceanic and Atmospheric Administration (NOAA) (2003).

145

5 A 4 0-8% SWD 9-11% SWD 12-14% SWD 15-17% SWD

Shoot flush rating

3

2

1

0 Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec

Month
5 B 4 0-21% SWD 22-31% SWD 32-50% SWD 51-60% SWD

Shoot flush rating

3

2

1

0 Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec

Month

Fig. 7-2. Shoot flush ratings of carambola trees in Krome very gravelly loam soil of South Florida in 2003 irrigated at four levels of soil water depletion (SWD) in an orchard (A, n = 51) and in containers (B, n = 22).

146

5 A 4 0-8% SWD 9-11% SWD 12-14% SWD 15-17% SWD

Shoot growth rating

3

2

1

0 Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec

Month
5 B 0 -21% SWD 22-31% SWD 32-50% SWD 51-60% SWD

Shoot growth rating

4

3

2

1

0 Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec

Month

Fig. 7-3. Shoot growth ratings of carambola trees in Krome very gravelly loam soil of South Florida in 2003 irrigated at four levels of soil water depletion (SWD) in an orchard (A, n = 51) and in containers (B, n = 22).

147

5 A 4 0-8% SWD 9-11% SWD 12-14% SWD 15-17% SWD

Flowering rating

3

2

1

0 Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec

Month
5 B 4 0-21% SWD 22-31% SWD 32-50% SWD 51-60% SWD

Flowering rating

3

2

1

0 Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec

Month

Fig. 7-4. Flowering ratings of carambola trees in Krome very gravelly loam soil of South Florida in 2003 irrigated at four levels of soil water depletion (SWD) in an orchard (A, n = 51) and in containers (B, n = 22).

148

5 A 4 0-8% SWD 9-11% SWD 12-14% SWD 15-17% SWD

Fruit load rating

3

2

1

0 Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec

Month
5 B 4 0-21% SWD 22-31% SWD 32-50% SWD 51-60% SWD

Fruit load rating

3

2

1

0 Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec

Month

Fig. 7-5. Fruit load rating of carambola trees in Krome very gravelly loam soil of South Florida in 2003 irrigated at four levels of soil water depletion (SWD) in an orchard (A, n = 51) and in containers (B, n = 22).

CHAPTER 8 SUMMARY AND CONCLUSIONS The objective of this research project was to determine the effects of irrigation at different soil water depletion (SWD) levels, on physiology, growth and yield of 2-yearold, container-grown, and 8-year-old, field-grown 'Arkin' carambola trees and to relate SWD to physiology, growth, and yield of carambola. Soil water depletion treatments were determined by continuous measurement of soil water content with multisensor capacitance probes (EnviroSCAN). Measurements of soil water content using multisensor capacitance probes were compared to a neutron probe measurements and tensiometer readings. Calibration equations for multisensor capacitance probes (EnviroScan) and a neutron probe in Krome very gravelly loam soil were also determined. The calibration equations obtained for the capacitance probe was y = 0.011x + 0.5206; r2=0.98, and that for the neutron probe was y = 29.05x-6.4; r2=0.95. In Krome very gravelly loam soil, water retention curves of tension determined with tensiometers vs. volumetric water content using capacitance probes or a neutron probe were fitted with van Genuchten’s model (1980). The r2 values indicated that the tensiometer tension readings were more highly correlated with neutron probe measurements of volumetric soil water content than with multisensor capacitance probe measurements. This suggests that capacitance probe measurements of soil water content were more variable in response to changes in soil texture throughout the soil profile than neutron probe measurements. The use of tensiometers was limited to a soil tension of 20 kPa due to air entry into the water column of the tensiometer and water column discharge. The use of a neutron probe by growers is 149

150 not practical because its radioactive source requires health and safety monitoring and a radiation license. Soil water content determined continuously with multisensor capacitance probes and computer software designed for irrigation management may be a practical method for irrigation scheduling in tropical fruit tree orchards. An increase in SWD resulted in reductions in ΨS, A, E, and gs of carambola trees grown in containers in Krome very gravelly loam soil. In an orchard, SWD never reached a point where there was a negative impact on ΨS, A, E, or gs. For trees in containers in Krome very gravelly loam soil, SWD levels above 50% caused a reduction in ΨS reduced gs. This resulted in a linear reduction in E and a sharp decline in A when gs fell below 50 mmol·m-2·s-1. Leaf gas exchange was more closely correlated with ΨS than with SWD level. Therefore, ΨS may be better basis than soil water content for irrigation scheduling for carambola in calcareous soils. There was no effect of SWD treatments on shoot growth of carambola trees grown in an orchard or in containers. In the orchard, sufficient soil water content from precipitation and the shallow water table possibly resulted in sufficient soil water content to obtain adequate vegetative growth and yields. For trees grown in containers, where lateral water movement and capillary rise were prevented, trunk diameter was decreased in trees irrigated at high SWD levels. Soil water depletion of 51-60% reduced total dry weight of trees in containers. Tree root:shoot ratios were not significantly different among SWD treatments because there were no significant differences in leaf and root dry weights among treatments. Fruit length, fruit number, and fruit weight did not differ significantly among SWD treatments in an orchard. Total soluble solids content and fresh

151 and dry weights of fruit were not related to SWD treatments in an orchard or in containers. Phenological cycles including shoot flush, extension shoot growth, flowering, and fruiting of carambola trees showed little response to irrigation at four SWD levels. This lack of response was likely caused by sufficient soil water due to precipitation and capillary rise from the high water table located at about 1-2 m below the soil surface of the orchard. Further studies over longer time periods are needed to better understand the phenology of these trees. Nonetheless, information gathered in this study on phenology of carambola may provide useful guidelines for orchard management in South Florida.

APPENDIX SOIL WATER DEPLETION, PHYSIOLOGY, GROWTH, AND YIELD, OF CARAMBOLA TREES IN KROME SOIL Table A-1. Effect of soil water depletion (SWD) on total number and weight of carambola trees grown in containers. Treatments Totalz fruit no. Totalz fruit wt. (kg) 0-21% SWD 29.17y a 2.26 a 22-31% SWD 20.33 a 1.66 a 32-50% SWD 24.33 a 1.54 a 51-60% SWD 34.17 a 2.18 a z Total of two harvests per tree (2002 and 2003), excluding immature fruit. y Means within columns followed by the same letter are not significantly different by Duncan’s multiple range test, P ≤ 0.05.

152

153

Fig. A-1. Soil water depletion treatments (SWD) as determined from measurements of soil water content using multisensor capacitance probes (EnviroSCAN) in an 8-year-old carambola orchard in Krome soil in South Florida. Field capacity was determined based on the pattern of soil water depletion over a period of 7 d. Visual stress symptoms (VSS) occurred when the leaves became wilted and chlorotic (photo inset in top right corner of graph).

154 Table A-2. Average soil water depletion (SWD) prior to irrigation of carambola trees in containers in the field and in an orchard in 2002 and 2003. Container average SWD (%) Treatment 2002
0-21 SWD 22-31 SWD 32-50 SWD 51-60 SWD 17 30 47 59

Orchard average SWD (%) Treatment
0-8 SWD 9-11 SWD 12-14 SWD 15-17 SWD

2003
20 31 50 59

Average
19 30 49 59

2002
9 10 14 17

2003
8 11 14 16

Average
8 10 14 17

155

2.4 2.2 2.0 1.8 1.6 1.4 1.2 1.0 0.8 0.6 0.4 0.2 0.0 Greenhouse Orchard

VP (kPa)

Location

Fig.. A-2. Vapor pressure (VP) difference between saturated and ambient air vapor pressure in the greenhouse and the orchard.

156

ΨS = - 0.57 MPa

A = 7.48 µmol m-2 s-1

Soil water content (%)

Field capacity

Onset of water stress

Calendar date

Fig. A-3. Soil water depletion (SWD) as measured with capacitance probes (EnviroSCAN) in an 8-year-old carambola orchard in Krome soils. Points of ‘field capacity’ and ‘onset of water stress’ were determined based on the pattern of soil water depletion. Measurements of stem water potential (ΨS) and net CO2 assimilation (A) indicated by the vertical double-line on the x-axis were chosen randomly at a point beyond the theoretical ‘onset of water stress’ point.

157

5

4

Shoot flush rating Shoot growth rating
Flowering rating Fruit load rating

3

2

1

0

4

3

2

1

0

4

3

2

1

0

4

3

2

1

0 Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec

Months

Fig. A-4. Ratings of the phenological cycles of carambola trees in Krome very gravelly loam soil in an orchard in South Florida in 2003 Trees were irrigated at four levels of soil water depletion (SWD) in an orchard (n = 51).

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BIOGRAPHICAL SKETCH Rashid Al-Yahyai was born in Al-Hamra, Oman, on February 8, 1971. He obtained his high school diploma in 1989 from Majid bin Khamis Secondary School at Al-Hamra, Oman. After graduation, he was enrolled in the College of Agricultural and Marine Sciences at Sultan Qaboos University, Oman. He earned his bachelor’s degree in May 1993 with a major in lant cience. Rashid earned a Master of Science degree in omology from the Department of Horticulture at Cornell University, Ithaca, New York, in May 1998. In January 2000, he was enrolled in the Department of Horticultural Sciences at the University of Florida. He conducted his research at the Tropical Research and Education Center, Homestead, Florida. He will be holding a faculty position at the Department of Crop Sciences, College of Agricultural and Marine Sciences at Sultan Qaboos University in Oman.

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