CRP 211

Applied Physiologies for Horticultural Crops

1+1

Course Teacher: Dr. K. Subburamu Theory Growth differentiation- Development definitions- types pattern of growth determinate indeterminate allometric growth. Growth Parameter analysis NAR RGR CGR harvest index partitioning efficiency. Growth regulators classification role of PGRs in horticultural crops. PGRs application in fruits vegetables. PGRs application in Spices and plantation crops flowers. PGRs for propagation. Senescence mechanism of senescence control of senescence by PGRs. Role of PGRs in Post harvest physiology. Nutrio Physiology importance of Macro Micro elements-essentiality mode of translocation. Concept of mobility of elements in plants indicator plants sampling for nutrient analysis. Deficiency of nutrients symptom expression general correction methods. Nutrient deficiency identification correction in fruits, vegetables, ornamentals. Nutrient deficiency identification correction in flowers, spices and plantation crops. Nutrient / PGRs formulation usage and precautions. Role of PGRs / nutrients for stress tolerance in horticultural crops. Effect of climate, soil, water and temperature on nutritional and physiological disorders in horticultural crops. Practical Measurement of growth - Estimation of leaf area leaf area index and Leaf Area Duration for different horticultural crops - Growth analysis Net Assimilation Rate Relative Growth Rate Crop Growth Rate - Study of Senescence, Senescence index and control measures - Seed and bud dormancy. Plant growth regulators commercial formulations role - rooting of cuttings - flowers, fruit set and development - ripening hormones - inhibitors / retardant / paclobutrazol. Nutrient Deficiency symptoms basic principles in identification -Deficiency symptoms in fruit crops, vegetable, flower crops Spices, plantation crops and correction measures - Rapid tissue test for nutrient

deficiency identification - Physiological disorders in horticultural crops. Visit to temperate / Subtropical regions for identification of nutrient deficiency and disorders in fruits flowers and vegetables. Lecture Schedule Theory 1. Growth differentiation- Development definitions- types pattern of growth determinate indeterminate allometric growth 2. Growth Parameter analysis NAR RGR CGR harvest index partitioning efficiency 3. Growth regulators classification role of Plant Growth Regulators in horticultural crops. 4. PGRs application in fruits and vegetables. 5. PGRs application in Spices and plantation crops and flowers. 6. PGRs for propagation. 7. Senescence mechanism of senescence control of senescence by PGRs. 8. Role of PGRs in Post harvest physiology. 9. Mid-Semester Examination 10. Nutrio-Physiology importance of Macro Micro elements-essentiality mode of translocation 11. Concept of mobility of elements in plants indicator plants sampling for nutrient analysis 12. Deficiency of nutrients symptom expression general correction methods 13. Nutrient deficiency identification correction in fruits, vegetables, ornamentals 14. Nutrient deficiency identification correction in flowers, spices and plantation crops. 15. Nutrient and PGRs formulation usage and precautions 16. Role of PGRs and nutrients for stress tolerance in horticultural crops 17. Effect of climate, soil, water and temperature on nutritional and physiological disorders in horticultural crops

Practical 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12. 13. 14. 15. 16. 17. References 1. Amar Singh, 1987. Fruit Physiology and Production 2. Bose, T. K., Mitra, S. K. and M. K. Sandhu, Mineral nutrition of fruit crops 3. Frank, B. 1995. Plant Pathology. 4. Kumar. 1995. Introduction to plant physiology. 5. Mallik, C. P. 1990. Textbook of plant physiology. 6. Mitcheli. 1988. Physiology of crop plants. Iowa State University. 7. Noggle and Fritz. 1983. Introductory to plant physiology. 8. Pandey, P. N. and B. K. Sinha. 1989. Plant Physiology. 9. Weston, C.P. 1994. Crop Physiology. 10. Williams P. Jacob. 1979. Plant Hormones and Plant development Measurement of growth components, metric analysis Estimation of leaf area leaf area index and LAD for different horticultural crops Growth analytical parameters measurement of NAR, PGR, CGR and HI Study of Senescence, Senescence index and control measures Preparation of commercial formulations and use of PGRs Role of PGRs on of rooting of cuttings Study of inhibitors / retardant / Paclobutrazol on horticultural crops. Study of PGRs for flowers, fruit set, development and ripening. Nutrient Deficiency symptoms basic principles in identification Deficiency symptoms in fruit crops and correction measures Deficiency symptoms in vegetable, flower crops and correction measures Deficiency symptoms in Spices, plantation crops and correction measures Visit to temperate / Subtropical regions for identification of nutrient deficiency and disorders in fruits, flowers and vegetables. Study of seed and bud dormancy Study of physiological disorders in horticultural crops Rapid tissue test for nutrient deficiency identification. Practical Examination

Chapter I Growth and Development Growth An irreversible increase in size, changes in form, shape. Evaluated by measuring of mass, length, height, area and volume. Restricted to living cells accompanied by metabolic process involving

synthesis of macromolecules, DNA, RNA, proteins, lipids, carbohydrates, secondary metabolites, growth regulators at the expense of metabolic energy. Differentiation Process involved in the establishment of localized differences in

biochemical and metabolic activity (specialized functions) and a structural organization that result in new patterns of growth. Development The term is used to encompass the activities resulting from growth and During development an orderly progression of transformations resulting in

differentiation. a characteristic form and chemical composition of plants takes place. Totipotency Isolated plant cells have the inherent capacity to give rise to a whole plant The ability of every living plant cell to produce the entire plant is called

totipotency Morphogenesis - Development of form or shape of cells and organs - Direction of cell expansion (decided by orientation of cellulose deposition) Control of cell division

Measurement of Growth etc. Vegetative growth Genetical makeup, Nutrients (16 elements), Environment Hormonal regulation, Tropic movement, Apical dominance Reproductive growth Genetical makeup Light Temp Moisture Hormonal regulation - External & Internal Nutrients Primary meristem - C: N - Ratio Apical meristem are formed during embryo development on the seed forms Secondary meristem The vascular cambium and meristematic zones are found near root and shoot tips in the vascular cambium. Determinate growth Germination Leaves flowers fruits Photoperiodism Thermoperiodism Vernalization Plant height, leaf size (length, width, area), fresh weight, dry weight of Cell number in tissues, organs. Conc. of chemicals constituent DNA, RNA, soluble protein, CH2O, lipids

different organs (stem, gist, leaves and fruits).

the growth of particular organ is determined.

Indeterminate growth

Root tips, shoot tips, apical meristem cells have capacity to regenerate

Determinate plants -

Apical bud ends with inflorescence - Monocarpic senescence

Indeterminate plants-

Apical buds always vegetative - Floral buds in axils Polycarpic senescence

Allometric Growth Growth of one part of plant may be closely related to growth of another part of the same plant, even though individual rates are different. Linear Curvilinear Linear Log y Slope b Log a 0 Log x 0 Log a Log x y = axb y= ax(b+c log x) Curvilinear Log y log y = log a + b log x

Phases of growth - Sigmoid Curve Y Steady state

Log Phase (exponential growth)

Initial lag phase

0
Growth analysis LA LAI LAD (in days) cm2 plant-1

X
S Shape or Wine cup shape

leaf area of plant / ground area occupied. L1 + (L1+1) 2 x t2 t1 L1 LAI at 1st stage L1+1 LAI at 2nd stage t1 & t2 time interval Leaf area ---------------------------Leaf dry wt. Leaf dry wt. ---------------------------Leaf area 1 = ------------SLW 1 = ------------SLA

SLA = cm2 g-1 SLW = mg cm-2 LAR = cm2 g-1

Leaf area ---------------------------Dry weight of the plant (DMA) W2 - W1 --------------------t2 t1 (Log e L2 Log e L1) x ----------------------(L2 - L1)

ULR or NAR mg cm-2 day-1

RGR = mg g-1 day-1

(Log e W2 Log e W1) ---------------------------(t2 - t1) W2 - W1 -------------------------P (t2 t1)

CGR = g m-2 day-1 P =

W1 & W2 plant dry wt. t1 & t2 - time interval

ground area on which W1& W2 has been estimated.

Vegetative growth Roots Food storage roots Propagative roots Pneumatopores Aerial roots Contractile roots Buttress roots Fungus roots Root nodules Stem Rhizomes Runner Stolons Tubers Bulbs Corms Thorns Tendrils - turmeric, zinger - strawberry - tubers at tip of stolons (Irish potato) - potato - fleshy leaves with a small stem at the lower end (onion) - gladiolus, corm, tuber rose - cacti, orchids - modified stem (Opuntia) - grapes (modified stem) - peas & cucumber (modified leaves) Chapter II Nutrio-Physiology Arnon & Stout 1939 - Criteria for essentiality of an element without which the plant cant complete its life cycle it cant be substituted by other element directly involved in plant metabolism either as a constituent of an essential metabolite or involved in biochemical reaction Beneficial elements CHO Na, Si, Co, V - carrot, beet, turnip, radish, - root suckers cherries, apple, pears - Avicenia (mangrove) - orchids, vanilla, large prop roots of banyan, corn - pull the plant deeper in to the soil (lily bulbs) - look like trunk gives stability to tree (fig) - Mycorrhizae of forest trees - Rhizobium Water storage roots - cucurbits

Cladophylls / cladodes / phylloclades

Ca Mg S Zn, Fe, Mn Cu, Mo, B Cl General Role of Mineral Nutrients

NPK

Toxic elements Al, Cd, Cr, Pb, Se, Ni, Br, I, F,

Nutritive function: Carbohydrate, Protein & lipids Structural frame work: By maintaining turgidity Structural Function: Cellulose, Hemicellulose, Pectin, Structural protein & lipids Balancing Function: Electrical Neutrality By K+ Cl Catalytic function: Metal activator / Cofactor in enzymes Specific Role of Mineral Nutrients Nitrogen: DNA, RNA, Enzymes, Amino acids, Proteins, Chlorophyll, Alkaloids, Cyanides Phosphorus: DNA, RNA, ATP, TPP, NADP, Phospholipids, Potassium: Stomatal movement, osmo-regulation, Cell turgidity, Structural frame work, Electrical neutrality, cell wall & membrane permeability, transport of assimilates Calcium: Cell wall, spindle formation in cell division, cell wall & membrane permeability, starch metabolism, calcium binding protein (Calcium calmodulin) Magnesium: Chlorophyll, Cofactor in enzymes involving PO4 transfer reactions (PO4Ase, kinase), cell wall & membrane permeability, Sulphur: Amino acids (Methionine, cystine, cysteine), Ferredoxin (non heam Fe-S compound) Enzymes tertiary structure, ~CoA, thiamine, diallyl disulpide. Mustard oil (glycosides, glucosinolates) Iron: Redox potential of the cell, Cytochrome. Plastoquinone, Ferredoxin, Enzymes (Dehydrogenase, catalase, peroxidase, oxidases) Zinc: Auxin metabolism, Tryptophan amino acid synthesis, Cofactor in enzymes

(alcoholic dehydrogenase, carbonic unhydrase) Manganese: Photolysis of water, Cofactor in enzymes (carboxylase & Decarboxylase), respiratory enzymes Copper: Plastocyanin, plant defense phenolic compounds, Cofactor in enzymes (Polyphenol oxidase, Laccase) Boron: Sugar transport, pollen viability Molybdenum: Cofactor in enzymes (NRAse), nitrogen metabolism Nitrogenase (Fe & Mo bearing N2 fixing microbes) Chlorine: Stomatal movement, Osmo-regulation, Cell turgidity, Electrical neutrality Beneficial elements Na Atriplex, halogeton, sugar beet, celery, spinach, turnip, coconut, cabbage, radish, rape, chenopodiaceae Si Rice Co Legumes Deficiency Symptoms: Nitrogen: Deficiency - starts from older leaves, whole plant affected Leaves small, dark green light green yellow brown die Plants - weak, small, few branches/tillers with anthocyanin Pigments purple colouration in leaves, petiole & stem, low soluble protein early maturity, short vegetative growth, Stem - weak, slender, spindly, Roots less branching Flowers few, poor fruit setting, small fruits, poor quality Phosphorus: Deficiency - starts from older leaves, whole plant affected Leaves small, thick, dark green with purple colouration Grey - brown die Plants Short, small, with pigmentation

Flowers delayed flowering - poor fruit setting, small fruits, poor quality, slow maturity Root Poor root formation

Potassium:

Deficiency - starts from margin/ tip of older leaves Leaves Yellowing starts from tip/leaf margin, bluish green leaves marginal, Scorching- tip burning, necrotic leaves die scorched to necrotic, leaf lets curl downward(curl towards under surface/ backward) Stem - weak Flowers few, poor fruit setting, small fruits with sour taste

Magnesium: Deficiency - starts from older leaves, interveinal chlorosis Reddening of leaves, mottling, reddish brown necrotic spots Leaves with marbling with tints of orange & purple colours. Interveinal chlorosis Zinc: Deficiency Rosettes terminal, starts from older leaves, interveinal chlorosis Leaves small, internodes constricted, mottle leaf Plants dwarf, rosette, malformed Flower flower shedding, poor fruit set, Fruits small, poor quality Calcium: Deficiency - starts from younger leaves, distorted, margin Irregular, Yellowing starts from tip of young leaves, leaves spotted & necrotic Terminal buds- twisted, hooked, die-back of young buds Root rotting Affect growing meristems

plants highly branched

Boron:

Deficiency - starts from younger leaves, chlorotic spots Yellowing starts from base of young leaves Leaves thickening, brittleness(transpiration affected), curling, wrinkling, wilting, root tips necrotic, Terminal buds- abnormal, twisted, hooked, die-back of young Buds stimulate lateral growth Young shoots discoloration, darkish blue colour, corky, cracked water soaked, blackening of vascular bundles in meristems plants highly branched pollen growth affected, (parthenocarpic fruit set) reduced flower & fruit formation

Sulphur:

General yellowing, similar to nitrogen deficiency Yellowing starts from younger leaves, Plants rigid, brittle, thin stem, shoot more affected than root

Iron:

Deficiency - starts from younger leaves Young leaves bleaching of Chlorophyll Green yellow white die

Manganese: Deficiency - starts from younger leaves, interveinal chlorosis Leaves- chlorotic, mottling, dark and light green bands Necrotic spots, streaks. Copper: Deficiency - starts from younger leaves, interveinal chlorosis

Leaves- chlorotic, withering of young shoots (lack of turgor), die back Gummosis (gum pockets surround the central pith), leaf curl upwards, poor quality of fruits Molybdenum: causes deficiency of nitrogen, interveinal chlorosis, whole plant affected, necrotic leaf margins due to NO3 accumulation, leaf lamina not formed, mid rib is present Indicator Plants N Maize and non legume cover crop in orchards P Tomato, maize K Potato, tomato, cucurbits, bean. alfalfa, clover, tobacco, corn and cotton Ca - Alfalfa Mg Portulaca, oxalis, apple, potato and cotton S Tea Fe Eucalyptus, acacia and morning glory Zn Citrus, peaches, field bean, onion, tomato, cotton and sorghum Mn Apple, cherry, citrus, sugar beet, raspberry and oats Cu Labiatae (Mint family-basil) and Caryophyllaceae (Pink family) Mo Brassica, beetroot, tomato, spinach, lettuce B Alfalfa, Apple, Pear, Cabbage, Brussels sprouts, & cauliflower General Deficiency Diseases N Firing old leaves Cucumber fruits pointed at blossom end, Onion small bulbs, early maturity P Delayed maturity, Dark green leaves K Tip burning, Marginal scorching, Firing of leaf tips and margins, apple marginal scorching, fruits small & poorly coloured tomato & grapes uneven ripening Potato tuber flesh is bluish

Mg Reddening, marbling in many crops S Tea Tea yellow Cruciferae narrow leaves Onion low pungency & more palatable Fe Bleaching of chlorophyll (devoid of chlorophyll) Ca Root rotting, Die back, Distorted, twisted and coiled Tomato & pepper Blossom-end rot Grapes Blossom-end rot & Stalk necrosis Pomegranate Fruit cracking splitting Apple Bitter pit, water core, Pear Cork spot Celery Black heart Brussels sprout Internal browning Carrot Cavity spot Chinese cabbage Tip burn Zn Interveinal chlorosis, Rosette, little leaf, dwarfening Apple small fruits Mn Dark and light green band of chlorosis, Grey speck Peas March spot/ Marshy spot Sugar beet- Speckled yellow Sugar cane Pahala blight Cu Gummosis, Withering (lack of normal turgor), Die back, Cupping or bowing up of leaves (pendula), splitting of young fruits Avocado S shaped shoots Citrus Gummosis or Exanthema (gummy excrescence in fruit rind & young stem) Onion Scales pale yellow Pear Witches broom Mo Citrus Yellow spot Cauliflower and Broccoli Whip tail Oats Blue chaff B Grapes Hen & Chicken

Apple Cork disease, Witches broom Cabbage & Brussels sprouts Hollow stem Turnip & Celery - Cracked stem, hollow & cracked root Cauliflower Brown or red rot Tobacco Top sickness Beet root Crown & Heart rot Pomegranate Fruit cracking splitting Amla Fruit necrosis Beans witches broom Physiological Disorders: Mango Black tip fumes of brick kiln Leaf scorch- Chlorine toxicity Spongy tissue Ca involved Soft nose or Tip pulp Ca involved Malformation nutrients? Mites? Viruses? Fungi? Citrus Granulation juice sacs hard (High temp., RH nutrients Zn Cu K involved) Rind pitting K reduced pitting Sapota Malformation of terminal shoot Fungi? Pomegranate Sudden decline Sudden drop in temp. late autumn and early winter Fruit cracking - Boron Annonas Stone fruit Competition among the developing fruits Fruit cracking Sudden high fluctuation in water supply Leaf scorching Chlorine toxicity Tomato Fruit cracking Sudden high fluctuation in water supply

Vegetable & Fruit crops Flower shedding biotic & abiotic stresses, Fruit drop - biotic & abiotic stresses Poor fruit quality Extreme temp. Poor flower colour High temp. Chapter II Plant Growth Regulators Hormone An endogenous compound, which is synthesized at one site and transported to another site where it exerts a physiological effect in very low concentration. But ethylene (gaseous nature), exert a physiological effect only at a near a site where it is synthesized. Classified definition of a hormone does not apply to ethylene. Plant growth regulators Defined as organic compounds other than nutrients, that affects the

physiological processes of growth and development in plants when applied in low concentrations. Defined as either natural or synthetic compounds that are applied directly to a target plant to alter its life processes or its structure to improve quality, increase yields, or facilitate harvesting. Plant Hormone When correctly used, is restricted to naturally occurring plant substances, there fall into five classes. Auxin, Gibberellins, Cytokinin, inhibitors and gas ethylene. Plant growth regulator includes synthetic compounds as well as naturally occurring hormones. Plant Growth Hormone The primary site of action of plant growth hormones at the molecular level remains unresolved.

Reasons Each hormone produces a great variety of physiological responses. Several of these responses to different hormones frequently are similar. The response of a plant or a plant part to plant growth regulators may vary with the variety of the plant. Even a single variety may respond differently depending on its age, environmental conditions and physiological state of development (especially its natural hormone content) and state of nutrition. There are always exceptions for a general rule suggesting the action of a specific growth regulator on plants. There are several proposed modes of action in each class of plant hormone, with substantial arguments for and against each mode. Hormone groups Auxins Substances generally resembles IAA and have the ability to stimulate the elongation of coleoptiles. Gibberellins are diterpenoids, which have the ability to elongate the stem of green seedlings especially certain dwarf and rosette types. Cytokinin Usually substituted Adenines, which resembles zeatin (Naturally occurring cytokinin in Zea mays) and have the ability to stimulate cytokinesis in cultures of tobacco cells. Ethylene Gaseous regulator that stimulate isodiametric growth in the apices of dicot seedlings. Inhibitors are regulators of growth, which originally depress the cell enlargement activity. Biosynthetic Precursor 1. IAA (Weak acid) Tryptophan

2. GA3 (Weak acid) 3. Ethylene 4. Cytokinin (Zeatin) 5. ABA (weak acid) Auxin Indole group IAA, IBA, IPA Naphthalene group - NAA Phenoxy group 2,4-D, 2,3,5 - T

Mevalonate Methionine Adenine Mevalonate Mevalonate

Dichlorophenoxy acetic acid (2,4-D) 2,4,5 Tricholorophenoxy acetic acid (2,4,5-T) IAA functions Promote the accumulation at K+ and ClIncreases the osmotic pressure Antioxidant Increase the elasticity of the cell wall and plasticity Reverses red light inhibition of mesocotyl elongation Geotropism IAA is transported to darker side of shoot Phototropism IAA is transported to dark side of shoot Leaf senescence delays Leaf abscission leaf application inhibit. Proximal application promotes Flowering may promote Fruit setting parthenocarpic fruit setting.

GA Gibberella fujikuroi (Fusarium moniliforma) Short tip, bud primordia, developing seeds GA antagonistic to ABA 55 Gas GA increases maleness, breaks dormancy, stem elongation, cell elongation in apical meristem. GAs increase the hydrolytic enzymes

Reduces the Juvenile period Cause flowering in plants belonging to diverse response types Delays senescence (retains chlorophyll) GA induces amylase protease, RNAse, Esterase, Acid PO4 ase Induce flowering and fruit set Parthenocarpic fruit set GA is a substitute for light and low temperature (cold) Cytokinins Substituted pureness Induces cell division Biosynthesis via RNA and biosynthesis of free cytokinins Breaks dormancy, causes femaleness Auxin and Kinetin ratio decides the root, short formation Root tip in the sit of cytokinin synthesis Structural component of RNA molecules Helps in protein synthesis. Delays senescence & protein degradation Counteract apical dominance Interact with light in seed germination, pigment formation and cell development Richmond Lang effect delay of senescence (1957) ABA Chloroplast could be the site of ABA synthesis Fruits constitute the riches source of ABA 2-trans ABA biologically active 2 cis ABA active 1. Synthesis by isoprenoid pathway from mevalonic acid 2. oxidation of xanthophylls. Mevalonic acid is also precursor of GA GA diterpene ABA sesquiterpene

ABA is antagonistic to GA Causes bud dormancy inhibition of germination Accelerate senescence stimulate abscission Causes closer of stomata Production of protein in mediated through ABA ABA is an inhibitor inhibits cell division and elongation ABA inhibits the synthesis of DNA and RNA Inhibit flowering in long day plants in SD condition Ethylene - antagonist CO2 All the plant part will produce ethylene It is regularly flushed out the plant Inhibition growth Leaf epinasty Abscission induction Fruit ripening Seed germination Bud sprouting CO2 fruit remain fresh Removal of CO2 by KOH enhances ethylene activity Increase later flow, Accelerate senescence Ethylene promote leaf abscission Increases the activity of IAA oxidase Increases the activity of chlorophyllase and causes breakdown of chlorophyll Waxing fruits results in low oxygen availability for ethylene production. Reversed by CO2 Chlorophyll degradation causes femaleness promotes abscission and senescence of flowers.

Ethylene Auto catalytic (or) ethylene production is common and ripening fruits and senescing tissues. Breaking of seed and bud dormancy Root initiation

Stem strengthening Lateral branching Leaf epinasty Flower initiation Flower and fruit thinning Fruit growth stimulation Initiation of fruit ripening Flower senescence Fruit and leaf degreening Leaf abscission and senescence

Phenols Monophenols Dipenols Polyphenols Salicylic acid, ferulic acid, P-coumaric acid Caffeic acid, catechol, , hydroquinone, colour of petals occur as glycosides anthocyanidin Anthocyanin Tannins gallic acid Lignin polymerised poly propane units Triacontanol long chained alcohol from alfalfa Increase dry weight, effect on photosynthesis Might increase uptake of nutrients Brassinolide steroid from pollen of the rape plant accelerate plant growth. Chloromequat (CCC) - reduce the size of the overall plant. Mepiquat chloride improves the standing power of crops. Salicylic acid Improves vascularisation Paclobutrazol Induces flowering CCC, Paclobutrazol Anti Gibberellins MH Anti auxin Alar B-Nine, Daminozide Dwarfening plant size

Uses of Plant growth regulators in Horticultural crops Germination and dormancy GA is the most potent germination promoter breaking seed dormancy Peach, sour orange, rough lemon, sweet orange, beans, peas, cabbage, cucumber, cucurbits, GA treated lettuce seed germinate in darkness. GA replaces cold requirements MH suppress sprouting in potatoes and onions GA breaks dormancy of potato tubers.

Rooting and Plant Propagation IBA -the best and most commonly used chemical IBA decomposed relatively slowly by the auxin destroying enzyme systems. IBA moves very slowly in the plant, much of it is retained near the site of application. produce strong fibrous root system NAA in more toxic than IBA. Amides of both compounds are also effective amide of NAA in less toxic than the acid itself 2,4-D and 2,4,5-T Promote root wig at low concentration but their toxic limit in near the opt. Concentration. Phenoxy acids produce bushy, stunted and thickened root system IBA- practical uses Rose , tea, winged bean, grapes, ficus, ixora, bougainvillea, jasmine, eggplant , hibiscus, apples, peach, plum, rubber Quick dip method: Basal end of cuttings dipped for a few second in a cove solution (upto 10,000 ppm in alcohol) Prolonged soaking method:

Basal end of cuttings are soaked in dilute solution (10-500 ppm) upto 24 hours. Powder method: Basal end of cuttings are treated with growth regulator in an earlier (clay or talc) conc. 500 to 1000ppm. Fruit set and Development -Auxin type compound most frequently used (NAA) GA set fruit in some sp that does not respond to others. Ethephon - female flowers in cucumber 2,4-D, NAA and IBA - cashew GA4 + GA7 and b- benzyl adenine- apple fruit quality Gametocides sterility Ethrel MH GA3 -

Cucurbits, Sugar beet grape, pepper, tomato cabbage, cauliflower, kale, lettuce grape, tomato,

TIBA Plant and Organ size GA breaks dwarfness

Increase size of grapes Triacontanol Brassinolide Increase dry weight, effect on photosynthesis Might increase uptake of nutrients steroid from pollen of the rape plant accelerate plant growth. reduce the size of the overall plant. Chloromequat -

Mepiquat and ethephon improves the standing crop.

Abscission The control of abscission does not reside in any one hormone or environmental factor but is regulated by a complex interaction of environment, hormones, and physiological status of plant Auxin, ethylene and ABA are most directly involved, others have indirect effect

Citrus Thinning Cycloheximides antifungal, antibiotic, - loosen citrus fruits (0.1/b per acre) it damages the flower and immature fruits. Release - effective abscission inducer. Apple thinning Ethrel Antitranspirant should be long lasting, cheap and Non toxic (PMA) Phenylmercuric acetate have toxic effects on the mesophyll and photosynthesis in addition to the influence on guard cells. Chemicals as metabolic antitranspirants ABA - bean, citrus, cucumber, pepper, tomato Chlormequot Daminozide 8 hydroxyquinoline IAA Phenyl mercuric acetate tomato tomato tomato, strawberry Tomato tomato, plant growth regulation (PGA)

Chapter III Senescence

Assimilate Partitioning in Horticultural crops

Transport system: Soil to root absorption Root to leaves Ascent of sap Leaves to atmosphere Transpiration Xylem transport Passive (water, minerals & cytokinin) Xylem = parenchyma + fibres + vessels + trachieds + ray cells Non living but functional (mainly dead cells) Unidirectional movement Phloem transport Active (photo assimilates- sugars, amino acids, organic acids, hormones & other biomolecules) Living functional Bi-directional movement Phloem transport - symplast via plasmodesmata depends on the rate of cyclosis within the cell. Provide a route for the movement of molecules from cell to cell without crossing the plasma membrane. Movement between cells in phloem sieve tubes occurs through sieve plates. Apoplast: movement of molecules through the (aqueous part of the cell wall matrix) apparent free space present in cell wall & intercellular airspace

cuticles, casparian strip and lignified walls are resistant to apoplastic transport. Phloem loading-Assimilate moves from mesophyll to sieve tube (Apoplast) Phloem unloading- Assimilate moves from sieve tube to sink cell (Apoplast) Apoplast pathway through cell wall & intercellular airspace Source regions of photo-assimilate production, export photo-assimilates Sink chlorophyllous tissues (leaves, stipules, fruit wall, young stem, pedicel, awns, peduncle, calyx, bract etc) - regions of photo-assimilate consumption, import photo-assimilates Growing regions, storage organs (respiration, growth and storage) Source strength: size x activity Differences in CO2 fixation (Rubisco & PEPCase) Leaf characters size, thickness, mesophyll size, compaction, vascular bundle Carrying capacity of sieve element (temp., H2O, nutrients, hormone) Sink strength - Potential capacity of the sink to accumulate assimilates Competition among different sink Harvest Index: HI = {Yield (Eco)/ Yield (Biol)} x 100 Harvest Increment: Photo-assimilates partitioned to harvestable organ - It is very difficult to estimate Improve HI in Annual horticultural crops Increase biomass production (DMA) & High photosynthetic rate Synchronized development of reproductive organ - Reproductively determinate High source strength at the time of sink differentiation Reduced growth of non harvestable organ Reduced leaf growth at reproductive stage with high LAD Optimum LAI and early peak LAI Prolonged and faster storage, enhanced competitiveness among of the storage organ Improved HI by increased size and number of sink organ

Decline in duration of Veg. growth and increased duration of Rep. growth. Reduced root weight with adequate nutrient and H2O Developmental plasticity (small surplus source for stress) Quick export of photoassimilates to avoid end product inhibition Efficient root system Limitations Source: leguminous vegetables Sink: Grain amaranthus Sink limitation: Late anthesis (Long Veg. phase) Indeterminate (Veg. & Rep. growth) Veg. growth at Rep. phase Less sink number and size Hormonal imbalance Stress Source limitation: Low canopy photosynthesis Low optimum LAI Slow peak LAI (lag vegetative growth) Low LAD at filling Early leaf senescence Stress nutrients, water PGRs: Auxin promotes source uptake ABA in leaves causes closer of stomata (Inhibit CO2 fixation) Cytokinin delays senescence of source and sink Cytokinin in sink increases photo assimilates import Ethylene induces senescence process. Assimilate Partitioning: Leaf subtending the inflorescence supply major portion of photo-assimilates for fruits development

In case of damage to the leaf subtending the inflorescence, the developing fruits mobilize assimilates from near by leaves Distance between the source and sink decides the rate of fruit development Current photosynthesis is important for fruit development in annual horticultural crops Different kinds of biotic and abiotic stresses adversely affect assimilate partitioning In perennials, stem always act as a strong storage pool for assimilates and nutrients. Previous season growth is important for higher productivity (In mango, SW monsoon rainfall is important for higher productivity)

Remobilizations occurs from stem to developing fruits

Fruit set, development & ripening

Flowering GA - Maleness ABA inhibit flowering in LDP and induce flowering in SD Plants CCC inhibit flowering in SD plants act as antigibberllins C2H4 induces femaleness in cucurbits at 3 leaf stage. C2H2, C2H4 and NAA induces flowering in pineapple Stress - Moderate stress early flowering Severe stress - delayed flowering - flower and pod shedding Parthenocarpy formation of fruit without seed i. ii. iii. Fruit development without pollination (Tomatoes, pumpkins, cucumber, bananas, pineapple) Fruit development due to stimulus received by pollination followed by unsuccessful fertilization (Poa) Abortion of embryos before attaining maturity of fruits (Cherries, peaches & grapes) Limitations of fruit set: Due to limited pollination Due to limited nutrients Due to abscission of flowers and young fruits

Pollenizer for cross fertilization for self barren trees honey bee for pollination parthenocarpic fruit set defective flowers (due to fundamental constituent of protoplasm) - imperfect flowers, heterostyly, dichogamy, due to fundamental constituent of protoplasm

Unfruitfulness Nutritive condition, soil type, water supply, cultural practice, wind low temp Pistillate flowers produce fruits parthenocarpically - Abortion of flower and flower buds before blossoming - Nonviable pollen & self sterility Self-sterility poor pollen tube growth in the style hormones, low temp - Difference in maturity of pollen grain & pistil (or) ovule - Delayed pollen germination Detective pistils in exhausted (or) weakened plants, overbearing, poor nutrition, drought Environment condition at the onset and during flowering affect fruit setting Nutrient supply hen & children disease in grapes Pruning and grafting alter apical dominance, control reproductive phase C/N ratio, normal bisexual flowers Locality soil, temp, humidity & light influence Season Age & vigour of plant, temp, light, H2O relations, rain wind proper stage of maturity at picking ensures better quality of fruits Fruit development: Fruit development depends on events taking place in ovule Fruits ripened ovary with attached parts 1. Well developed carpel Orange, grape 2. Epigynous flower receptacle in a part of the fruit (eg.) apple, banana, cucumber 3. Ripened ovary tomato Berry Entire ovary wall develops into a fresh tomato, brinjal, guava, grapes, avocado Hesperidium Citrus multicellular enlarged, juicy, saclike out growth from the surface of the carpal walls are edible

Pepo One or three celled syncarpous pistil with parietal placentation Drupe Exocarp thin, mesocarp fleshly, endocarp stone almond, mango, peach, apricot, plum, coconut, cherry Pome Fleshy edible part is thalamus Aggregate Collection of simple fruits or fruit lets Strawberry edible fleshy and much elongated receptacle rasp berry. Multiple fruit inflorescence takes point in the formation of fruit - pineapple, mulberry, fig, jack fruit Pericarp Date, custard apple Pericarp & placenta tomato, brinjal Mesocarp papaya, mango, melon Mesocarp & endocarp banana Mesocarp, endocarp & placenta cucumber Thalamus strawberry, pineapple, apple Thalamus & pericarp guava Peduncle & cotyledon cashew Endosperm coconut Receptacle fig, pear Bract, perianth & seed jack Juicy placental hair citrus Juicy outer coat of seed pomegranate Phases of fruit growth I Fruit growth rates two important stages Growth of pericarp Epicarp, mesocarp & endocarp (Rapid and slow growth) Growth of embryo & endosperm Simple sigmoid- apple, pineapple, strawberry, pea tomato Double sigmoid peach, apricot, plum & cherry (stone fruits); fig & grapes (Nonstone fruits) II Mobilization: Large size fruits import food from other parts of the plant III Fruit size:

Fruit size correlated with cell size (cherries) Large fruit size through inter cellular spaces development in the second half of fruit enlargement (apple) Loading of sugars into the fruit seed regulate fruit growth Removal of fertilized ovule terminates fruit growth Geometry of fruits reflects seeds distribution Seeded regions of fruits distinct from parts of unseeded region Fruit size is + vely to seed number (strawberry) Seedless pear oval; seeded pear pear shaped Seed influence strong on the growth of fleshy fruits

IV Role of seed:

Parthenocarpic fruits differ in shape from seeded fruits of same variety

Growth substances Endogenous Auxin play important role Single sigmoid Auxin controlled Double sigmoid I phase Auxin and GA controlled II phase osmotic accumulation of carbohydrates ABA and C2H4 involved in fruit drop Tomato seeds contain ferulic acid which causes dormancy Fruit drop: Fruit resulting from self pollination are more susceptible to fruit drops Auxin content is low - fruit drop is common ABA and Ethylene involved in fruit drop C2H4 inducing compounds are successfully used in mechanical harvesting Pectin, starch, sugars, sugar-acid ratio Maturity attainment of full size Matured fruits maximum starch and sp. gr Late harvest Lowers shelf life Fruit Ripening: Ripening Loss of chlorophyll

Maturity age, shape, angularity, size, appearance, texture, sp. gr

Increase in brix Increase in titrable acidity Decline in pectin and water soluble pectin Lowering respiration increase shelf life Ripening: Fruit ageing - Softening of the flesh - Changes in pigmentation and flavour - Loss of chlorophyll - Appearance of carotenoids - Increase in sugars & volatile compounds adds flavour Polysaccharides sugars, in acids CO2 & C2H4 production increase Increase in activity of malic enzyme and pyruvic carboxylase leading to climacteric rise in CO2 production climacteric fruits hydrolytic enzymes During ripening sudden upsurge in respiration followed by deterioration banana, guava, mango, fig, jack, papaya rich in carotenoids fructose disappears require high energy for fruit ripening increase in RNA & mRNA Increase in chlorophyllase, hydrolase, lipase, lipoxidase, peroxidase C2H4 bring about climacteric in non-climacteric fruits Unsaturated hydrocarbon promote ripening Single gradual decline in respiration citrus (lemon, lime, oranges, grapes, pomegranate, pineapple rich in anthocyanins N2 & CO2 & O2 prevent the climacteric rise. Softening cell wall-degrading enzymes Hydrolysing of cell content (pectic enzymes) middle lamellae Increase in methylation of galacturonic acid Hydrolytic changes sugar formation Changes in pigments Decrease in chlorophyll (chlorophyllase) increase in carotenoids & anthocyanins (phytochrome involved) Low temperature Increase in colour & High temperature Poor colour development Flavour substances Esters, aldehydes, ketones, loss of astringent materials, decrease in

phenolics Climacteric fruits- banana, mango, apple, pear, tomato Increase in respiration during ripening-after reaching a peak the rate of respiration falls Low O2 and high CO2 affect climacteric rise Polythene storage lowers O2 and increase in CO2 extends shelf life C2H4 stimulates climacteric rise Non-climacteric fruits citrus, pepper, peanuts (Steady respiration) Ripening Respiration is a source of energy for synthesis of new enzymes required for ripening C2H4 ripening hormone, but it has no effect on ripening but de-greening occur

Photoperiodism (Vince-Prue, 1975)

Day length controls initiation and development of flowers Asexual reproduction Formation of storage organs Onset of dormancy Specific biochemical pathways regulate different responses

Garner & Allard Maryland Mammoth tobacco Borthwick & Parter Soybean Hamner & Bonner Xanthium strumarium Chailakhjan Chrysanthemum & Perilla Melchers & Lang Hyocyamus niger (henbane) Vince Prue specific biochemical pathways regulate different responses Night length is more important than day length SDP (< 12 hrs) Dahlias, Asters, Chrysanthemum, Goldenrod, poinsettias, Salvias, Straw berries, maize femaleness, sweet potato, Soybean var. Biloxi Chenopodium, coffea arabica SDP- I Qualitative (or) absolute Chrysanthemum, Soybean Cv. Biloxi Chenopodium, Ipomea batatas Coffee arabica, Xanthium strumarium (cocklebur)

Tobacco Cv Maryland mammoth II Quantitative cosmos, amaranthus, zinnia, sunflower, rice, potato, strawberry (runner LD), onion (bulb-LD), datura, dahlia, aster, poinsettia, LDP - (> 14 hrs) Beets, lettuce, spinach, Spinacea oleracea Spinach, Oats, barley, Opium poppy, Radish - Raphanus sativus, alfalfa, sugarcane, Lettuce LDP - I Absolute (or) Qualitative: Beta vulgaris, Avena sativa, Raphanus sativus, Spinacea oleraceae, Hordeum vulgure II Quantitative: Lettuce, alfalfa, opium (poppy), wheat, barley, oats, Brassica rapa, Pisum sativus, Secale cereale IDP DNP - (12 14 hrs) Indian grass & other grasses, Ocimum basilicum - (not regulated by day length) roses, sunflower, tomatoes, beans, carnations, nasturtium, snapdragon, Dandelions, calendulas, paddy, cucumber, maize Critical day length - expression of photoperiodic response SDP do not flower if the day length is longer than their critical day length. LDP do not flower in light periods shorter than their critical period. Hypothetical Plant hormone Florigen Floral stimuli is transmittable Photoperiod (hours) LLLLLL 8 h LLLLLL 8 h LLLLLL LLLLLLL 8 h LLLLLLLLLLLL 16 h DDDDDD LL SDP LDP D DDDDDDDDDDDD 16 h DDDD 8 h DDDD 8 h xxxxxxxxx LL DD X SDP LDP SDP LDP F F V F F V Duration of dark without any break is more important for perceiving photo stimuli. and lower night time temperature. DDDDD L DDDD 16 h LLLLLL 16 h DDDDD 8 h LLLLLLLLLLLLLLL 16 h SDP F V V V V F LLLLLL DD LDP V F F F F V LLLLLL

DDDDDDDDDDDD 16 h DDDDDD

Thermoperiodism - The beneficial effects on plant growth of alternating either day-time

Vernalisation - The enhancement of subsequent flowering response by low temp (0100C) treatment of inhibited (or) young seedlings Gassner, Munreek and Whyte, Lysenko, Gregory and Purvis, Lang and Melcher, Requirements: water, oxygen and low temp. 0-10 C (opt. 3 - 7 C) Seed Vernalisation Chicory - Cichorium intybus Carrot Daucus carota Sugar beet Beta vulgaris Winter cereals Oats, Barley, Rye, Wheat, Indian Mustard, Turnip, Lettuce

Plant Vernalisation - Rape, Celery, Digitalis, Oenothera Hypothetical plant hormone - Vernalin