INTRODUCTION
Start flower induction and differentiation
Seed development : a sequence of events controlled by the genotype Maturation: set of successive stages in preparation for successful seed germination Flowering and pollination are not uniform in the same plant or within a population 2
INTRODUCTION
Initial studies: aimed at determining phenological differences among species and the optimum harvest time
First proposed criteria to identify seed maturity: - Time from planting or seedling emergence to harvesting - Seed moisture content and morphological characteristics to identify seed maturity
SEED DEVELOPMENT
Delouche (1971) : Seed maturation is a process that comprises a set of morphological, physical, physiological and biochemical events that occur from ovule fertilization to the moment in which seeds become physiologically independent of the parent plant
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HISTODIFFERENTIATION
1960s and 1970s: individual seeds x plant population Seed sampling at pre-defined intervals and identification of parameters associated with maturation progress Definition of seed changes in seed technology research
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SEED MOISTURE CONTENT Ovule fertilization and evolution during maturation Dry fruits x Fleshy fruits
Final dehydration
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Fresh weight
Moisture content
Dry weight TIME Variations in moisture content, fresh weight and dry weight during maturation of seed produced in dry (- - -) (or fleshy fruits ( ). 8 (Carvalho and Nakagawa, 2000)
SEED SIZE
R5
R6
SEED SIZE
R6
R7
R8
10
Reference Carvalho (1972) Jacinto and Carvalho (1974) Nagai (1973) Carvalho and Yanai (1976)
Days after flowering necessary for seeds of different species to attain maximum size during maturation (Carvalho and Nakagawa, 2000) (Carvalho 11
DRY WEIGHT
number of cells Number of cells/seed x 10-6 dry weight
6 30 4 20 2 10 (Dry weight (mg/seed) 40
Egli
11
12
Dry weight
Egli
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Species Cotton Peanut Oat Field bean Maize Soybean Sorghum Wheat
Reference Carvalho (1972) Carvalho et al. (1976) Frey et al. (1958) Neubern and Carvalho (1976) Hunter et al. (1991) Andrews (1966) Kersting et al. (1961) Carvalho and Yanai, 1976
Seed moisture content of different species at the time they reach reach maximum dry weight during development 14
GERMINATION
Primary root protrusion x Normal seedlings Germination x Dormancy during maturation
VIGOR
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Initial germination (days) after ovule fertilization in some cultivated cultivated species (Delouche, 1971) (Delouche, 16
Moisture content (%) 55.3 50.6 48.1 39.8 27.2 14.1 11.0 8.8 9.9 10.7
Seed dry weight (mg/seed) 20.40 25.34 28.80 34.10 36.44 34.10 37.24 37.54 37.90 37.84
Germination (%) 05 28 77 37 11 09 06 81 83 65
Vigor (mg dry weight / seedling) 5.0 4.07 7.47 7.69 4.86 4.88 5.53 9.57 10.72 9.34
Variation of seed moisture content, dry weight, germination and vigor during wheat maturation, cv. IAS -54. (Carvalho and Yanai, 1976) 17
CONCEPTS a) Seed maturity is identified by the maximum dry matter accumulation b) Seed physiological maturity is reached when there are no significant increases in seed dry weight c) Seed physiological maturity occurs when seeds reach maximum dry weight, germination, and vigor 18
PREVAILING IDEA
Seed physiological maturity = maximum dry weight Relative Maturity, Morphological Maturity, Mass Maturity, Harvest Maturity, Agronomic Maturity, Time of Harvest
Determination of physiological maturity in individuals or in plant communities
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MOISTURE CONTENT
SIZE
VIGOR
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Moisture content (%) 72.9 60.5 56.8 43.9 38.2 21.8 41.0 30.0 18.1
Seed dry weight (mg/seed) 61.4 158.4 171.0 202.8 203.8 194.8 208.6 206.4 194.8
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Mean values of moisture content and dry weight during maturation of bean seeds, cv Carioca (Neubern and Carvalho, 1976)
Difficult to identify the exact point of seed physiological maturity (maximum dry weight) NEED TO INCREASE PRECISION: - Number of statistical replicates - Reduce harvesting intervals - Additional care in weighing Use of 14 C to monitor reserve accumulation
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PM
100 80
Percentage
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63
77
90
Days after R3
Marcos-Filho (1979) 24
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Black Layer
Mature
Maturation of tomato
Physiological potential Overripe
Contreras
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Species Lettuce Onion Tobacco Maize Inbred Single hybrid Double hybrid Soybean Tomato Wheat
Average and range in days before or after physiological maturity and seed vigor maturity of different cultivated species (TeKrony and Egli, 1997) () (-): days before seed maximum dry weight ((+): days after seed maximum dry weight
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Germination() FD 00 00 65 54 SD 68 68 -93
Soluble Sugars() FD 1.1 1.5 2.2 1.6 SD 0.1 0.2 0.2 0.4
Percentage germination, leakage of sugar and protein from soybean seeds, cv soybean Chippewea, submitted to drying at different maturation stages (Adams and Rinne, 1983) () FD seed threshing and fast drying immediately after harvest; SL slow seed drying still attached to pods. () Germination (%); Seed leachate (mg/seed)
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Dry weight
Physiological Potential
Maturation of tomato
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Effect of harvest time on maize seed yield in two locations of Parana state, Brazil (Hadlich, 1983) 31
RH RH
RH
RH
RH
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FIELD WEATHERING
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PM 94 85 89 83 71
HT 88 83 85 28 62
14* 58 85 66 13 40
28* 55 41 57 07 05
Vigor (accelerated aging) of Kent soybean seeds as influenced by harvest time for five experimental years (TeKrony et al., 1980).
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- Harvesting at physiological maturity? - Variations in seed moisture content and in plant characteristics - Visual parameters: black layer, milk line, seed and/or fruit color, glumes and pedicel
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- Uniformity of maturation
Soybean
Tomato
Cotton
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II
III
Carrot
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Umbel / Plant 1 11 24
11 58 31
75 65 54
54 43 37
Number of umbels per plant, contribution to plant seed production (%), germination (%), vigor (germination first count - %) and weight of 1000 seeds associated with umbel order in carrot (Nascimento, 1991) 38
- Monocots
endosperm; Dicots
cotyledons
- Final yield: seed number + seed growth rate + duration of seed fill
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RESERVE ACCUMULATION DURING SEED DEVELOPMENT 1. NUTRIENT TRANSLOCATION FROM THE PARENT PLANT
Photoassimilates (sugars, amino acids and other solutes) are transported via phloem Plant reserve accumulation in endospermic seeds Nutrient translocation to seeds Short distance transport
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starchy endosperm
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RESERVE ACCUMULATION DURING SEED DEVELOPMENT 1. NUTRIENT TRANSLOCATION FROM THE PARENT PLANT
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f ee
h
ct
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Mature seeds two or three main types of reserves Synthesis is parallel during seed development Synthesis occurs in different cellular compartments: starch, in amyloplasts lipids, in spherosomes proteins, in cytosol and endoplasmic reticulum
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2. NUTRIENT ASSIMILATION
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MAIZE
Dry Weight 300 mg 200 DNA 0,06 0,04 mg mg
100
0,02
30 Proteins mg 20 RNA
0,5 0,3
10
0,1
15
30
30
45
46
embryo
whole seed
2. NUTRIENT ASSIMILATION
LEGUMES
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50 40 30 20
Proteins
Percentage
Lipids
Carbohydrates 10
30
50
70
Changes in carbohydrate, lipid and protein contents during soybean seed development (Adapted from Konno, 1979)
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cell division
140
cell elongation
1400 7
100
RNA proteins
1000
60
600
20
200
10
22
26
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proteins (mg/cotyledon)
DNA (g/cotyledon)
Stage R4 R5 R6 R7 R8
Dry weight
Proteins Oil Sugars (mg/seed) (mg/seed) (mg/seed) ---------2.5 42.0 71.9 73.4 0.1 26.7 31.7 36.2 1.3 19.3 30.8 32.4
HORMONES
Effects of water deficits Water excess Time of occurrence Association with temperature stress
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Association of water deficit and thermal stress during soybean seed filling (Frana Neto and Krzyzanowski) (Fran 53
APROSMAT, 2006
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- Temperature Critical value: 35oC during seed filling Forced maturation and greenish seeds
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Germination (%)
Color
Emergence (%)
Initial
Yellow Green
3
months
Initial 75 52
3
months
Initial 87 58
3
months
88 60
87 19
73 20
88 38
- Light Photosynthesis x plant and seed performance Retention of flowers and fruits
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Seed 4-stage developmental pattern: cell division, cell elongation, reserve accumulation, desiccation Seeds do not germinate while attached to the parent plant During most of seed development: Formation and activity of enzymes involved in synthesis processes, managed by m-RNA
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Metabolic reversion from synthesis to hydrolysis Synthesis and activity of ABA High concentration during embryogenesis Synthesis and activity of ABA Decrease during late maturation desiccation Management by m-RNA
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HORMONE INFLUENCE
cyt
gib
aux
ABA
Fertilization
Maturity
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synthesis
mobilization
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Desiccation during seed maturation: developmental pattern, driving seed embryo to a quiescent stage in preparation for germination Desiccation Tolerance: The ability to recover biological functions after drying to equilibrium with moderately dry air and then resume normal function when rehydrated This ability depends on the ability to maintain membrane structure and prevent protein denaturation
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Desiccation Tolerance: Intolerant Phase: Cell division and elongation + part of reserve accumulation period Tolerant Phase: Final phase of reserve accumulation Premature and rapid desiccation: Damage to enzyme and protein synthesis Loss of cell turgidity, damage to cell membranes, enzymes, proteins and nucleic acid structure
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- LEA (late embryogenesis abundant) Proteins - Antioxidant systems - Soluble sugars: raffinose, stachyose, sucrose - Heat shock proteins - Slow drying
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