Evaluation of Four Treatments to Break Seed Dormancy

of Sunflower Inbreds

A Research Study Presented for the

Master of Science in
Agriculture and Natural Resources Degree
University of Tennessee at Martin

Submitted by: Rogelio Marchetti

December 2012
 ii 
 
ABSTRACT

Seed dormancy can be a major problem for seed companies. In sunflower (Helianthus

Annuus, L.) breeding programs, dormancy limits the number of crop cycles per year; it also leads

to asynchronous blossom times, restricting the opportunity to make crosses between plants. This

study evaluated simple techniques to break dormancy in sunflower. A preliminary laboratory

study confirmed that only 32% of non-treated plants emerged, whereas exposure of seed to a

cold treatment improved the germination to 92%. The subsequent field experiment tested four

treatments intended to break possible dormancy effects. Six sunflower inbred lines were grown

and harvested 30 days after flowering to assure enough time for embryo development while

promoting dormancy. The first two treatments were aimed at removing physical dormancy by

chipping a minuscule section of the pericarp or by exposing the seed to temperature oscillation to

degrade impermeable layers. The other two treatments worked at the hormone level of dormancy

by placing seeds in a saturated atmosphere of ethylene gas or by submerging the seed in ethrel

(liquid ethylene). The four treatments were compared to a control, in which seeds were kept in

the cold at night and at room temperature during day. The experiment was arranged in a split plot

randomized complete block design (RCBD) and evaluated with analysis of variance. The highest

mean emergence was obtained in the control (54%) and temperature oscillation (52%)

treatments. Mean emergence for the seed chipping treatment was 32%, which was significantly

(P<0.05) lower than all other treatments. However, there was a significant (P=0.0062) interaction

between treatments and lines. Although some of the treatments increased germination in some

lines, they also depressed germination in other inbreds. No treatment improved germination

consistently over the control while most of them added complexity to seed handling.

iii 
 
 Table of Contents

Page

Chapter I. – Introduction .............................................................................................................1

Economic importance ...................................................................................................................1

Objective .....................................................................................................................................4

Chapter II. – Literature Review ...................................................................................................5

Seed germination ..........................................................................................................................5

Sunflower .....................................................................................................................................7

Dormancy in sunflower seeds ......................................................................................................9

Hormones ...................................................................................................................................10

Breaking seed dormancy ............................................................................................................11

Chapter III. – Materials and Methods .......................................................................................13

Site description ...........................................................................................................................13

Material selection .......................................................................................................................13

Experimental treatments .............................................................................................................15

Experimental design ...................................................................................................................16

Data collection............................................................................................................................17

Statistical analysis ......................................................................................................................17

Chapter IV. – Results and Discussion ........................................................................................18

Chapter V. – Conclusions ............................................................................................................20

Literature Cited ...........................................................................................................................21

iv 
 
 List of Tables

Page

Table 1. Sunflower production by country. .................................................................................... 2

Table 2. Germination of six sunflower inbreds using warm and cold germination tests.............. 14

Table 3 List of treatments tested for breaking dormancy in sunflower inbreds. .......................... 15

Table 4. ANOVA table for percent emerged plants of dormant Sunflower inbreds. ................... 18

Table 5. Germination percentage for each seed treatment ............................................................ 18

Table 6. Germination percentage by seed treatment and sunflower line ...................................... 19

v 
 
 List of Figures

Page

Figure 1. Sunflower production areas. ............................................................................................ 2

Figure 2. The three phases and physiological processes of germination ........................................ 7

Figure 3. Seed dormancy and the control of germination ............................................................. 11

Figure 4. Layout of research plots in the field using a split plot randomized complete block
design. ............................................................................................................................ 16
 

vi 
 
 CHAPTER I – Introduction

Sunflower (Helianthus annuus L.) is an annual plant native to North America that

belongs to the Compositae (Asteraceae) family. This family is distinct in that the inflorescence is

formed by hundreds of flowers acting as one. The slim and long flowers that encircle the

sunflower inflorescence, called “ray flowers”, are sterile and their main purpose is to attract

insects. The disk florets, in the center of the inflorescence contain fertile tubular flowers that will

produce achenes, the main interest of this crop. Wild sunflowers are an open pollinated and self

incompatible species; thus wind and insects play a vital role in the reproductive process.

Breeding techniques improved self compatibility in commercial hybrids; however, the presence

of insects is required for seed production (USDA, 2000).

Economic Importance

Due to the high oil content, sunflower seed is primarily crushed and consumed as one of

the most important edible oils. High oleic hybrids that produce seed containing more than 80%

oleic acid are becoming more desirable because they provide a healthier option for human

consumption, placing sunflower into the economically generous specialty sector (Putnam et al.,

1990).

Besides oil, sunflower has many other uses; confectionary hybrids are utilized in the food

industry, whereas bird seeds and sunflower pellets are part of the feed industry. Sunflower is also

finding new markets and can be used to produce biodiesel (Putnam et al., 1990).

1 
 
 Wide adaptation of sunflower allows it to be planted worldwide (USDA, 2000). Due to

low costs of production, good market price and robust performance, it is usually grown in dry

areas with poor soil conditions, limiting the true potential yield of this crop (Figure 1).

Russia is the top producer of sunflower seed, followed closely by Europe and Ukraine

(Table 1). Sunflower production is distributed among many countries compared to other crops,

where two to three countries grow a big portion of the global volume.

Figure 1. Sunflower production areas (www.wikipedia.com).

Table 1. Sunflower production by country – FAO, 2010 (www.fao.com).

Rank Country 106 M Tons

1 Russia 7.4
2 EU 6.9
3 Ukraine 6.3
4 Argentina 3.6
World Total 33.4
 

2 
 
 Throughout history, natural selection, along with selection by humans, shaped sunflower

to its present-day phenotype. Current hybrids are quite different from wild sunflowers; they are

single headed, short plants with a short pollination period (USDA, 2000). The major impacts on

crop improvement and yield of sunflower have been made through plant breeding techniques.

Agronomic traits including disease, insect and herbicide resistance and soil and longitude

adaptation have been added to commercial hybrids through plant breeding.

Whereas plant breeders of other crops, such as soybeans and maize, utilize the latest

techniques in biotechnology making faster improvements with transgenes, sunflower breeders

face a different reality. Potential drift of transgenic events to wild sunflowers and consumer

reluctance to accept genetically modified foods thwart the possibility of utilizing biotechnology

in sunflower (N.S.A., 2000). Today, sunflower breeders rely only on conventional techniques; it

may take up to 15 years to develop and release a new hybrid. To reduce this time frame, most

seed companies implement “winter nurseries,” which are off-season locations that enable

breeders to complete up to three crop cycles in one year, reducing the total number of years to

develop a new inbred to four. Although the benefits of using winter nurseries are worthwhile,

some challenges arise. At harvest the seed is not fully developed and when trying to minimize

days between harvest and planting, germination problems may affect the subsequent planting. A

study on wild sunflower shows that seed maturation influences germination, and seeds are most

mature at 21 days after flowering (Seiler, 1993). Late and uneven emergence are signs of

dormancy issues. If plants do not develop at the same pace, pollination synchrony becomes a

problem, reducing the possibility of making crosses.

3 
 
Objective

The main purpose of this study was to evaluate economical and simple techniques to

remove or reduce dormancy effects in seeds of sunflower inbreds. Reducing dormancy will

enable breeders to:

 Replant nurseries faster to achieve three crop cycles per year.

 Obtain homogeneous emergence.

 Promote uniform parent - progeny blossom times to enable crossing among inbreds.

4 
 
 CHAPTER II – Literature Review

Seed Germination

Crop establishment is the first step and one of the most important stages in the vegetative

cycle of crop plants. Uneven emergence and low stand density will compromise the subsequent

stages of growth, limiting potential yield and economic outcome. Growers work to provide the

right conditions for planting: preparing a proper seedbed (moisture and texture), selecting an

appropriate variety, planting at the right time and depth, using a seed treatment, etc.

Unfortunately there are aspects over which farmers have no control and they must rely on the

seeds to emerge rapidly and evenly from the soil.

Nonogaki et al. (2010) notes that germination starts with water imbibition followed by

physiological changes in the seed and is complete with the appearance of the radicle through the

seed structures. The authors explain germination through three phases:

Phase I:

Due to a negative water potential the seed rapidly absorbs water until the cells are

completely hydrated. This crucial step re-activates the seed from a hibernation state with

physiological and chemical changes. Probably the most important change is the resumption of

energy metabolism. Mitochondria and enzymes increase the rate of respiration providing energy

to sustain this rapid growth. Structures and organelles that were damaged during the drying and

re-hydration processes are repaired and replaced. Residual mRNAs, which are formed during

seed development, survive desiccation and contribute to the fast restoration process after

imbibition. Later new mRNA’s are created to continue with protein synthesis (Nonogaki et al.,

2010).

5 
 
Phase II:

During this stage, water uptake decreases, forming a plateau (Fig. 2). Cells are now turgid

and hormones take over the process. The most influential hormones in seed germination are

abscisic acid (ABA) and gibberellic acid (GA). ABA is utilized as a messenger within a plant to

suspend growth. ABA is involved in leaf abscission and also in preventing germination when

environmental conditions are not adequate. On the other side, GA has positive effects on

germination; it is a precursor of alpha-amylase, an enzyme that mobilizes seed reserves. The

interaction or balance between these two antagonistic compounds is what triggers germination.

The presence of ABA deactivates GA and reduces GA biosynthesis while the opposite effect is

found when GA concentration is higher than ABA. Environmental factors such as temperature,

light, nutrition, and smoke regulate ABA:GA ratio, controlling germination.

During phase II, fully hydrated cells impose pressure on the soaked and surrounding weakened

structures of the seed, allowing the radicle to emerge. At this point, the second phase, as well as

germination, is completed (Nonogaki et al., 2010).

Phase III:

The last stage, also called postgermination, is the visible growth of the radicle. This

process is characterized by an increase in water uptake, cell division and reserve mobilization.

The seedling also grows, based on mitotic division and cell enlargement (Nonogaki et al., 2010).

6 
 
 Figure 2. The three phases and physiological processes of germination (Nonogaki et al. 2010)

Sunflower

The center of origin of sunflower is located in the central plains of North America (Harter

et al., 2004). This plant produces fruits called achenes, which are non-dehiscent fruits surrounded

by the pericarp (external hull), with the actual seed inside. Throughout history, wild sunflowers

have survived extreme weather conditions, diseases, and pests. Dormancy is a survival

technique, similar to hibernation in animals, which prevents seed from germinating until

favorable conditions are present.

Mercer et al. (2006) compared wild sunflower accessions from different parts of the

United States with pure sunflower inbreds. The researchers made various crosses between them

and studied the degree of dormancy of the progeny. Parents and hybrids were planted in the field

and results showed that wild-crop hybrids prevail over wild-wild crosses, giving higher

7 
 
percentage of germination (65% for wild-crop versus 34% for wild-wild) and less dormancy

(58% for wild-crop and 27% for wild-wild). Wild-wild hybrids struggled to survive and

remained dormant for longer periods. This evidence indicates that the parents of sunflower

hybrids have a strong influence in determining dormancy.

Dormancy can be biologically described as: “The absence of germination of an intact,

viable seed under germination favoring conditions with a specific time lapse” (Hilhorst, 1995).

There are different mechanisms and processes involved in seed dormancy. Baskin and Baskin

(2004) divided seed dormancy into five types:

1) Physiological dormancy (PD)

Physiological dormancy is the most common dormancy present in angiosperm and

temperate species, including Helianthus annuus (sunflower), Lycopersicon esculentum (tomato),

and Lactuca sativa (lettuce). Internal changes at the hormone level are needed for the seed to

germinate. This group varies from deep dormancy (3-4 months) to nondeep (less than one

month) (Baskin and Baskin, 2004).

2) Morphological dormancy (MD)

This group presents small but differentiated embryos that have no physiological

dormancy. They require more time for the embryo to mature in order to germinate. Example:

Apium graveolens (Baskin and Baskin, 2004).

3) Morphophysiological dormancy (MPD)

Seeds with morphophysiological dormancy are characterized by small and differentiated

embryos but also have physiological dormancy. Thus they need more time to grow plus a

8 
 
treatment to overcome dormancy such as stratification or GA treatment (Baskin and Baskin,

2004).

4) Physical dormancy (PY)

Physical dormancy is caused mainly by impermeable seed coats that prevent water

uptake. Thick and lignified cell walls are responsible for this physical barrier; water repellent

compounds (waxes, cutin and suberin) that cover the seed can also limit water uptake. Until the

external layers become permeable to water by temperature oscillation, by seed going through the

digestive system of animals, or by freezing and thawing cycles, the germination process will not

start (Baskin and Baskin, 2004).

5) Combination dormancy

Seeds in this group show a combination of physical dormancy accompanied by

physiological dormancy. To overcome dormancy, seeds need to break PY first, allowing water

permeability, and, once imbibed, the embryo will release PD after stratification (Baskin and

Baskin, 2004).

Dormancy in sunflower seeds

A study by Brunick (2007) at the University of Oregon showed that sunflower genotypes

presenting pericarp and coat dormancy are the most difficult to include in breeding programs due

to the complexity of the genes involved, whereas embryo dormancy tends to be more malleable.

In this study, the researchers demonstrated that the number of layers of seed coat plays a crucial

role in imposing extensive dormancy. Wild sunflower seeds tend to be sturdier, having more

layers, while commercial sunflower seeds allow water and oxygen to flow through the

9 
 
teguments, facilitating germination. Another important finding in this research is the response to

daylight. Although sunflower is not greatly affected by this environmental signal, Brunick (2007)

found that germination was greater in light compared to the dark. He recommends alternating 12

hours of light with 12 hours of dark to maximize germination.

Hormones

Hormones play a decisive role in seed dormancy. Finch-Savage and Leubner-Metzger

(2006) describe the relationship between the ABA and GA: germination is not an absolute

process that responds to the presence or absence of one or the other hormone. It is a proportion

or ratio balance where the predominance of one over the other creates a gradual change.

Figure 3 describes the effect of the environment on the ABA:GA ratio and ultimately on

germination (Finch-Savage and Leubner-Metzger, 2006). In the perception phase, external

factors such as light, temperature and water create signals to genes responsible for the synthesis

and degradation of ABA and GA. During the integration phase, the balance of ABA:GA

orchestrates the subsequent steps transferring into the response stage. When environmental

conditions promote ABA prevalence, this triggers more ABA synthesis and GA degradation,

causing the seed to stay dormant. Under favorable conditions, GA synthesis prevails, causing

ABA catabolism and germination initiation.

10 
 
 Figure 3. Seed dormancy and the control of germination (Finch-Savage
and Leubner-Metzger 2006)

Breaking seed dormancy

Just as there are different types of dormancies, there are different methods to overcome it.

For example, if a seed presents physiological dormancy, the application of exogenous hormones

or exposure to varying temperatures will remove it.

Water imbibition has many positive effects on removing dormancy. Maiti et al. (2006),

successfully utilized priming, where seeds are submerged in water for 12-24 hours, to effectively

break dormancy in sunflower seeds. Priming works first by activating the germination process.

Secondly, it works by washing away ABA and other compounds that have negative effects on

11 
 
germination. Lastly, imbibition weakens teguments and hard structures, thereby removing

physical dormancy.

Pallavi et al. (2010) utilized temperature treatments to break dormancy of sunflower

seeds. Temperature treatments of 60oC for 15 minutes significantly increased germination over

control by desiccating waxes, weakening the impermeable layer, and allowing water to be

absorbed.

Growth regulators have a positive effect on germination. Application of exogenous

compounds, such as GA or ethylene, will modify the ABA:GA ratio removing dormancy effects,

allowing seed to germinate. This was confirmed by Borghetti et al. (2002), when seeds

submerged in ethrel (25 ppm) had significantly increased germination. Seiler (1994) reported

that the use of 1 mM solution of gibberelic acid (GA3) doubled the germination rate of dormant

sunflower seeds

Bratcher et al. 1993, successfully utilized tetrazolium chloride (TTC) to improve

germination of wild sunflower seeds. Treatments included perforating the seeds with a needle

and submerging the seeds in TTC. These physical mechanisms, or scarification, debilitate the

external structures or teguments facilitating water uptake and oxygen inflow.

12 
 
 CHAPTER III – Materials and Methods

Site description

This study was conducted at the Pioneer Hi-Bred facility located in Woodland, California

(UTM Zone 10 coordinates: E: 600939, N: 4281280), which is the main summer location for

sunflower research. Summers are hot with an average temperature of 35.8oC and winters are cold

with average minimum temperature of 3.1oC. Precipitation occurs almost exclusively between

November and March, with an average of 470 mm per year (Western Regional Climate Center,

2011). The soil is classified as Yolo Silt Loam by the Soil Survey of Yolo County (Andrews,

1972). This soil is a member of the Fine-silty, mixed, superactive, nonacid, thermic Mollic

Xerofluvents.

Material Selection

Materials to be tested were carefully selected based on potential dormancy effects.

Pioneer has an annual evaluation on all inbred lines comparing relative maturity. Six lines with

the longest maturities over the last three years were used in this study. Three of them are

characterized as high oleic materials, which are also known to exhibit dormancy. The following

inbreds were evaluated: conventional line 1 (L1), conventional line 2 (L2), conventional line 3

(L3), high Oleic line 1 (L4), high Oleic line 2 (L5), high Oleic line 3 (L6)

Foundation seed was produced at the Wiamea station located in Kekaha, Kauai, HI. The

seed was planted on January 31st, 2012. To be consistent across different maturities, each line

was harvested exactly 30 days after flowering. Once harvested, lines were kept in a cold room

(48°F and 45% RH) until shipped to Woodland, CA on May 27th, 2012 for the main study.

13 
 
 To determine a baseline and identify if seeds were dormant, a germination test was

performed in the laboratory under controlled conditions. A warm germination test was compared

to a cold germination test to evaluate possible dormancy effects. In the warm germination test,

50 seeds of each inbred were placed in a germ towel and sprayed with water. The towel was

folded to assure contact between seeds and then placed in a growth chamber at 25°C for seven

days. The percentages of normal, dormant and dead seed were calculated. The cold germination

test was the same as the warm test, but required a pre-chill of seven days at 10°C prior to the

seven days at 25°C. Exposure to the pre-chill technique greatly improved the germination rate in

these inbreds (Table 2). This evaluation under controlled conditions clearly revealed the presence

of dormancy problems in all lines. Once dormancy was confirmed, the main study was

conducted by testing four proven methods of breaking dormancy.

Table 2. Germination of six sunflower inbreds using warm and cold germination tests

Line Germination under warm test Germination under cold test

Line 1 42% 92%
Line 2 39% 92%
Line 3 31% 92%
Line 4 34% 94%
Line 5 27% 94%
Line 6 37% 94%

14 
 
Experimental treatments

Seeds from each line were divided into five groups and exposed to the described

treatments using four replications (Table 3).

Temperature changes were used in the first treatment, in which seeds were exposed to 8

cycles of alternating temperatures: 30 minutes of -80°C followed by 30 minutes of room

temperature (20°C). These temperature changes promote minuscule fractures in the outside

layers of the seed, facilitating water imbibition and oxygen flow.

The second treatment was a mechanical cut on each seed to assure water uptake and

oxygen flow into the seed. Seed chipping was manually performed with dog nail clippers cutting

the tip of each hull without damaging the seed.

In the third treatment, seeds were enclosed in a chamber with a saturated atmosphere of

100% ethylene gas for 12 hours at 20°C; this treatment was intended to remove physiological

dormancy.

The fourth treatment was ethrel application in which seeds were submerged into a

solution of 30ppm ethrel (Ethepon 2SL diluted with water) for 15 minutes. This treatment

presoaks the seeds and triggers germination by increasing the GA concentration.

Table 3 List of treatments tested for breaking dormancy in sunflower inbreds.

Dormancy effect to act on Treatment Action

1- Physical limitation
2- Physical limitation
3- Physiological changes
4- Combination of 2 & 3
5- Control
Freezing – Thawing cycles
Seed chipping
Ethylene exposure (gas)
Ethrel submersion (liquid)
No treatment
+20°C to - 80°C
Mechanical cut on the hull
12hs @ saturated atmosphere
15 minutes

15 
 
 The last treatment in the experiment was the control, in which seeds were kept in the cold

room (9°C and 45% RH) during night and brought out during day (20°C) until planted. This is

the normal procedure the sunflower breeding program utilizes because seed needs to be weighted

and conditioned for packet filling.

Experimental design

A split plot randomized complete block design was utilized to establish the plots in the

field (Figure 4). There were six lines, and five dormancy treatments replicated four times

(blocks). Seed treatments were the main plots while the subplots consisted of the six sunflower

inbred lines.

20 Chip L2 L5 L4 L3 L6 L1
19 Submer. L1 L3 L5 L6 L4 L2

Rep 4
18 Control L4 L2 L6 L1 L5 L3
17 Temp L2 L6 L3 L4 L1 L5
16 Gas L3 L1 L2 L5 L6 L4
15 Submer. L2 L5 L4 L3 L6 L1
14 Gas L1 L3 L5 L6 L4 L2
13 Temp L4 L2 L6 L1 L5 L3 Rep 3
12 Control L2 L6 L3 L4 L1 L5
11 Chip L3 L1 L2 L5 L6 L4
10 Control L2 L5 L4 L3 L6 L1
9 Gas L1 L3 L5 L6 L4 L2
Rep 2

8 Chip L4 L2 L6 L1 L5 L3
7 Temp L2 L6 L3 L4 L1 L5
6 Submer. L3 L1 L2 L5 L6 L4
5 Temp L2 L5 L4 L3 L6 L1
4 Chip L1 L3 L5 L6 L4 L2
Rep 1

3 Gas L4 L2 L6 L1 L5 L3
2 Submer. L2 L6 L3 L4 L1 L5
1 Control L3 L1 L2 L5 L6 L4
1 2 3 4 5 6
Figure 4. Layout of research plots in the field using a split plot randomized
complete block design.

16 
 
 Before planting, the field was pre-irrigated with 25mm of water by drip irrigation and

sprayed with glyphosate to eliminate any emerging weeds. Soil temperature at planting depth

(2.5 cm) was 23°C. The experiment was planted on June 1st, 2012. The planter utilized was the

Almaco Seed Pro, which is specifically designed for research purposes and provides detailed

information such as planting depth, numbers of seeds planted and spacing. The average number

of seeds per plot was 50 and the spacing between plants was 17 cm.

Immediately after planting, 37mm of water was provided by drip irrigation to assure

water imbibition. Weather conditions were optimal through the entire germination process, with

an average day temperature of 28°C, with some highs of 38°C, and an average night temperature

of 19°C.

Data Collection

Observations were made between June 6th and June 27th, 2012. Number of emerged

plants per plot was recorded on a daily basis until 21 days after planting. After 21 days, very few

plants will emerge and flowering will be delayed, limiting the opportunity to make breeding

crosses. Germination was calculated as an absolute value; the plant was either emerged or not.

Fully expanded cotyledons was the threshold to determine that a plant had emerged.

Statistical Analysis

Data collected during the experiment were analyzed using Proc GLM in Statistical

Analysis Software (SAS). Data were analyzed as a split plot randomized complete block design.

The Ryan-Einot-Gabriel-Welsch multiple range test was utilized to determine if there were

significant differences among means.

17 
 
 CHAPTER IV – Results and Discussion

Based on the statistical analysis, there were significant (P < 0.0001) differences among

treatments (Table 4). Mean emergence of the control treatment was equivalent to emergence of

the temperature treatment and was significantly higher than the rest of the treatments (Table 5).

Mean germination was 54% for the control and 52% in seed that received the temperature

treatment. Seeds exposed to ethylene gas had a mean germination of 45%, which was the same

mean as those submerged in Ethrel. Seed chipping treatment had 32% germination, which was

significantly lower than all other treatments (Table 5).

Table 4. ANOVA table for percent emerged plants of dormant Sunflower inbreds.

Source of Variation Degrees of Sum of Mean Square F Value Pr > F

Freedom Squares
Block 3 1290.00 430.00 5.78 0.1001
Treatment 4 6953.80 1738.45 23.38 < 0.0001
Main Plot Error 12 892.33 74.36 0.37 0.9691
Inbred 5 13830.26 2766.05 13.88 < 0.0001
Treat * Inbred 20 8973.40 448.67 2.25 0.0062
Subplot Error 75 14949.66 199.32

Table 5. Germination percentage for each seed treatment.

Treatment Germination %
Control 54a †
Temperature 52ab
Gas exposure 45b
Submersion 45b
Chipping 32c
† Means followed by the same letter are not significantly different (P<0.05)
using Ryan-Einot-Gabriel-Welsh

18 
 
 The statistical analysis also indicated that there was a significant interaction (P = 0.0062)

between treatments and inbreds (Table 4). Seed chipping improved germination for Line 1 from

a mean of 50% to 63%, while the same treatment negatively affected germination for Line 6

from a mean of 61% to 30% (Table 6). Line 6 had the highest germination rate across treatments,

but germination fell noticeably with seed chipping. This was also confirmed by Brunick (2007)

where achene excisions produce mixed results and germination varied with genotypes. The

control was the only treatment that showed consistent emergence compared to other treatments.

Germination across lines was also variable. The expectation that linoleic lines present

better germination compared to high oleic lines could not be confirmed. Two of the three linoleic

lines had germination rates above or equal to 50%. Germination percentage in high oleic lines

ranged from 30-37% except for line 6, which had 61% germination, the highest value in the

study (Table 6). The results are similar to the findings of Murcia et al. (2002). However, the

small set of inbreds tested is not sufficient to draw conclusions on germination rates between oil

profiles.

Table 6. Germination percentage by seed treatment and sunflower line.

Seed Ethylene Ethrel
Temperature Control Mean by line
Chipping gas submersion
Line1 43 63 47 47 51 50
Line2 54 33 24 32 43 37
Line3 64 36 49 63 73 57
Line4 46 16 40 36 46 37
Line5 32 15 43 31 42 33
Line6 73 30 69 61 70 61
§
Mean by treatment 52  32 45 45 54
 §
For line means within treatments, LSD0.05 = 19.9

19 
 
 CHAPTER V – Conclusions

Although the average germination values in this study were low (30 to 60%), it is

important to note that the conditions imposed were skewed towards dormancy. Only late

materials, which require longer time to mature, were used in this study. These lines were

harvested 30 days after flowering, allowing just sufficient time to complete the physiological

cycle without drying naturally.

Based on the premises stated in the objectives and the inbred lines tested, none of the

seed treatments provided superior germination over the control. Also, the seed treatments require

additional steps and are more expensive.

The observed interaction between lines and treatments could be due to different types of

dormancies affecting each line. This interaction is a major drawback because it prevents the

utilization of a single treatment for all lines, adding complexity to the entire process.

20 
 
 Literature Cited

Andrews, W.F. 1972. Soil survey of Yolo County, California. USDA Soil Conservation Service,
Washington DC.

Baskin, J.M. and C. Baskin. 2004. A classification system for seed dormancy. Seed Science and
Research 14: 1-16.

Bratcher B. C., J. M. Dole, and J. C. Cole. 1993. Stratification improves seed germination of five
native wildflower species. Hortscience 28(9): 899-901.

Borghetti F, F. Nakamura Noda and C. Martins de Sá. 2002. Possible involvement of proteasome
activity in ethylene-induced germination of dormant sunflower embryos. Brazilian Journal of
Plant Physiology 14: 125-131.

Brunick R. 2007. Seed dormancy in domesticated and wild sunflowers (Helianthus annuus L.).
[online] Available at:
http://ir.library.oregonstate.edu/xmlui/bitstream/handle/1957/7865/Brunick%20Dissertation
%20%20Final%20PDF.pdf?sequence=1

Finch-Savage, W.E. and G. Leubner-Metzger. 2006. Seed dormancy and the control of
germination. New Phytologist 171: 501-523.

Food and Agricultural Organization (FAO). 2010. Agribusiness Handbook – Sunflower crude
and refined oils. [online] Available at:
http://www.fao.org/docrep/012/al375e/al375e.pdf
Harter, A., K. A. Gardner, D. Falush, D. L. Lentz, R. A. Bye, and L. H. Rieseberg. 2004. Origin
of extant domesticated sunflowers in eastern North America. Nature 430: 201–205.
Hilhorst, H.W. 1995. A critical update on seed dormancy. I. Primary dormancy. Seed Science
Research 5: 61-73.

Maiti, R.K., P. Vidyasagar, S. C. Shahapur and G. J. Seiler. 2006. Studies on genotypic

variability and seed dormancy in sunflower genotypes (Helianthus annuus L.) Indian J. Crop
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