Watermelon
Crop Information
Breeding Methods
- by Todd C. Wehner
- Department of Horticultural Science
- North Carolina State University
- Raleigh, NC 27695-7609
Breeding Objectives
Major objectives for watermelon breeding include proper
fruit type, early maturity, high fruit yield, high sugar
content, tough flexible rind, and proper seed type. It is
important to determine breeding objectives carefully before
starting variety development. For example, seed type changes
significantly for different market classes. Parental lines
for seedless hybrids should have small seeds, whereas confectionery
seed types should have large seeds. For commercial varieties,
black seeds are preferred because of their contrast with
red, yellow, or orange flesh. Also, white seeds indicate
immaturity to buyers, so white mature seed color can be
a confusing trait for them. Most of the old varieties are
diploid, open-pollinated or inbred lines, but hybrid diploid
and hybrid triploid varieties are taking over the commercial
market in the United States.
After determining the breeding objectives, methods for
measurement of the traits of interest should be developed,
selection methods should be determined (specifying the operations
to be carried out for each generation), and parents with
high expression of the traits of interest should be chosen.
Vine type should be long for commercial production and dwarf
(bush) for home garden. It may also be possible to use the
dwarf plant type for once-over harvest in commercial production.
Sex expression should be monoecious, with a ratio of 7 staminate:1
pistillate flowers, or better (preferably 4:1). Andromonoecious
sex expression and ratios of 15:1 are more typical of older
varieties.
For production in most areas of the United States, watermelon
must have resistance to fusarium wilt. Races 0 and 1 are
common, and race 2 is becoming important, especially in
Texas and Oklahoma where plastic mulch culture and fumigation
are less common. Production areas in the southern United
States usually have anthracnose race 1 and may also have
problems with race 2. Gummy stem blight is a disease for
which resistance is needed in most southern production areas.
Powdery mildew is becoming a problem, especially in the
western United States (possibly because of a new race),
and should be a breeding objective for new varieties. Bacterial
fruit blotch was a problem in the 1990s, and resistant accessions
have been identified. The disease can be effectively controlled
by genetic resistance and by large-scale seed testing followed
by destruction of contaminated seed lots. Protection from
viruses in the United States production areas should include
resistance to papaya ringspot virus-watermelon strain (formerly
watermelon mosaic virus-1), watermelon mosaic virus (formerly
watermelon mosaic virus-2), and zucchini yellow mosaic virus.
Finally, breeding objectives should emphasize early maturity,
high fruit yield, durability for shipping, high internal
quality, freedom from internal defects (hollowheart and
rind necrosis), and proper seed type in a diploid (seeded)
or triploid (seedless) hybrid. Internal quality traits include
dark red flesh, high sugar content, proper sugar to acid
ratio, excellent flavor, high nutritional value (vitamins
and lycopene), firm (not soft) and non-fibrous texture.
Seeds should be black color, medium size (or small for inbreds
to be made into tetraploids), and few to medium quantity
per fruit (few for consumers, but medium to keep seed costs
down). Flesh color should be dark red (Y gene with modifier
genes) with uniform color throughout the fruit. For specialty
types, flesh color could be bright orange (yo gene), canary
yellow (C gene), or white (Wf gene). Other colors such as
salmon yellow (y gene) exist (Table 3.2), but are not preferred
because the flesh looks overmature. Older varieties have
light red flesh, but dark red is becoming the preferred
type. Diploid inbreds should be made into tetraploid inbreds
and tested for fertility, seed yield, and ability to set
fruit using controlled pollination. Tetraploid lines for
use in triploid seedless hybrid production can be induced
with colchine. Finally, triploid hybrids should be tested
for absence of seed coats in the fruit within a range of
production environments.
Variety Development
There were no defined varieties of watermelon before the
1820s. Early varieties include ‘Black Spanish’
(imported to United States from Portugal in 1827), ‘Carolina’
(available at least since 1827), and ‘Imperial’,
‘Mountain Sprout’, ‘Seminole’, and
‘Mountain Sweet’ (introduced by southern growers
from 1840 to 1850). Other heirloom varieties include ‘Bradford’,
‘Clarendon’, ‘Odell’, ‘Ravenscroft’,
and ‘Souter’ (originating in South Carolina
before 1850). Classic watermelon varieties include ‘Peerless’
or ‘Ice Cream’ (1860), ‘Phinney Early’
(1870), and ‘Georgia Rattlesnake’ developed
by M.W. Johnson in Atlanta, Georgia about 1870.
Planned variety development programs began in the United
States in 1880 to 1900. Important varieties developed for
the southern United States included ‘Cuban Queen’
developed and marketed by Burpee in 1881, ‘Round Light
Icing’ (1885), ‘Kolb Gem’ developed by
Reuben Kolb of Alabama in 1885 and marketed by D.M. Ferry,
‘Florida Favorite’ selected from the cross of
‘Pierson’ x ‘Georgia Rattlesnake’
by Girardeau in Monticello, Florida in 1887, ‘Dark
Icing’ developed in 1888 by D.M. Ferry, and ‘Dixie’
selected from the cross of ‘Kolb Gem’ x ‘Cuban
Queen’ or ‘Mountain Sweet’ by George Collins
in North Carolina and marketed by Johnson and Stokes. Important
varieties developed for the western United States included
‘Chilean’ (black or white seeded) brought from
the west coast of South America and introduced to California
in 1900, ‘Angeleno’ developed by Johnson and
Musser in Los Angeles, California in 1908, and ‘Klondike
Solid’ and ‘Klondike Striped’ of unknown
origin developed about 1900. Important varieties developed
for shipping include ‘Tom Watson’ developed
by Alexander Seed Co. in Augusta, Georgia in 1906, and ‘Stone
Mountain’ developed by Hastings Co. in Atlanta, Georgia
in 1924.
Important varieties developed in the latter part of last
century have built on past accomplishments. ‘Charleston
Gray’ (USDA, Charleston, 1954), ‘Crimson Sweet’
(Kansas State University, 1963), ‘Calhoun Gray’
(Louisiana State University, 1965), and ‘Dixielee’
(1979), ‘Jubilee’ (1963), and ‘Smokylee’
(1971) (all from the University of Florida) have high resistance
to Fusarium wilt. ‘Dixlee’ (University of Florida,
1979) and ‘Sangria’ F1 (Novartis, 1985) have
dark red flesh. ‘Millionaire’ F1, 3x (Harris
Moran, 1992) and ‘Royal Jubilee’ F1 (Seminis)
have consistently high yields. ‘Crimson Sweet’
(Kansas State University, 1963) and ‘Sugarlee’
(University of Florida, 1981) have high soluble solids.
‘Kengarden’ (University of Kentucky, 1975) has
dwarf vines. ‘Tri-X-313’ F1 3x (Novartis, 1962)
is seedless. ‘Minilee’ (University of Florida,
1986), ‘Mickylee’ (University of Florida, 1986),
‘New Hampshire Midget’ (University of New Hampshire,
1951), ‘Sugar Baby’ (M. Hardin, Oklahoma, 1955),
and ‘Tiger Baby’ (Seminis) are icebox size.
‘Yellow Doll’ (Seminis, 1977) has canary yellow
flesh.
Breeding Plan. Once the breeder has determined
the objectives of the program, the choice of parental materials
is one of the most important aspects of a breeding program.
Using knowledge of the crop and predicting the traits consumers
will be interested in having in future varieties, the breeder
gathers parental lines for crossing. The breeder should
know which parent will contribute the traits of interest,
and which methods will be used to evaluate the progeny for
those traits. Thus, it is often necessary to collect and
evaluate large numbers of PI accessions, varieties, and
breeding lines for the traits of interest to identify appropriate
parents to use in the program. This work often continues
in parallel with the main part of the breeding program.
The next step is to determine the breeding method to use
for each part of the program. It is important for the breeder
to consider the advantages and disadvantages of particular
breeding methods, and how they can be incorporated into
the overall breeding plan. Also, it is common to use more
than one breeding method at a time in order to accomplish
several sets of objectives. For example, one part of the
program might be to use recurrent selection to develop a
base population with general adaptation and the proper fruit
type that also has high yield and early maturity. A second
part of the program might be to use pedigree selection on
the cross of two lines to develop inbred lines with the
high yield, early maturity, and proper fruit type of one
parent, and the dark red flesh color, high sugar content,
and firm crisp flesh texture of the other parent. A third
part of the program might be to use backcross breeding to
make a canary yellow flesh version of an elite red-fleshed
hybrid with top performance.
Recurrent Selection. Although watermelon
is a cross-pollinated crop, population improvement methods
popular in some cross-pollinated crops have not been used.
The main reason for that appears to be the large size of
the plants, and the low rate of natural outcrossing that
occurs. Also, because there are few plant breeders working
on watermelon, and because of the requirement for many qualitative
traits to be present in the new varieties being tested for
release, it is expensive to spend additional years in population
improvement for quantitative traits.
It may be possible to improve quantitative traits such as
yield in watermelon using recurrent selection i.e. repeated
selection and massing of selected plants, but the populations
should probably be developed initially to have the necessary
qualitative genes in them. Those would include proper flesh
color, fruit size, and disease resistance. Due to large
plant size and a 5-month generation time, recurrent selection
methods should be those that have few generations per cycle,
and few plants per family (or single-plant selection).
One approach would be to develop an elite population by
intercrossing two to four of the best red fleshed hybrids
available, trying to choose a set that was genetically unrelated.
A population with a wide genetic base could also be developed
by intercrossing 20 or more elite varieties by hand for
two or more generations, and using bees in an isolation
block for two or more generations before beginning a mild
selection pressure for important quantitative traits such
as yield. Simple recurrent selection (Fig. 3.8) could be
used for selection among single-plant hills for a set of
highly heritable traits. A more complex method such as reciprocal
recurrent selection would permit simultaneous improvement
of two populations for combining ability for yield (Fig.
3.9). This would be an expensive program to run, but would
produce two populations that could be used to develop inbreds
to be used as the female and male parents (respectively)
of elite hybrids.
During population development, it would be necessary to
identify methods for yield testing that were efficient for
use in large yield trials. The usual guidelines for recurrent
selection are to test at least 200 individuals (or progenies
of individuals) per population, and to select at least 20
to intercross for the next cycle of selection. A yield trial
involving 200 replicated families would require more resources
than many breeding programs could afford if the trial were
done using current methods.
Recurrent selection could be used to improve quantitative
traits, such as yield, which are difficult to improve using
qualitative methods such as pedigree and backcross breeding.
Each year, the improved population would be used to begin
the development of inbred lines to feed into other parts
of the breeding program.
Pedigree Breeding. Probably the most common
method for watermelon breeding is pedigree. In pedigree
breeding, the breeder begins by choosing two or more adapted
parents, which complement each other in their traits. For
example, one parent might be generally good (yield, earliness,
type) except for disease resistance and the other might
be generally good (yield, earliness, type) except for fruit
quality. The objective would be to produce new lines with
high yield, early maturity, proper type, high fruit quality,
and good disease resistance. The varieties or breeding lines
are crossed to form the hybrid (F1) generation, which is
then self- or sib-pollinated to form a segregating (F2)
population (Fig. 3.10). The F2 is self- or sib-pollinated
while selecting for traits having high heritability to form
the F3 generation. If multiple plants are tested from each
selected F2 plant, then the breeder concentrates on selecting
the best plants in each of the best F3 families. This might
include selection in the seedling stage in the greenhouse
in the F2 and F3 generations for disease resistance such
as fusarium wilt races 0, 1, and 2 and anthracnose races
1 and 2.
Beginning at the F4 generation, selection would begin to
emphasize family-row performance for quantitative traits.
Plants within family-rows that have excellent performance
for qualitative traits should be selected for the next generation.
As the families reach six generations of self-pollination
(S6 or F5), they become more uniform, and can then be handled
as inbred lines. This could include selection using eight-plant
plots for early flowering, number of pistillate flowers,
and fruit number. The number handled might decrease from
54 F2 plants of a cross to 36 F3 families, 24 F4 families,
and 18 F5 lines.
Single-seed-descent is a modification of pedigree breeding
in which inbred lines are developed rapidly by self-pollination
in greenhouses and winter nurseries, and selection is not
practiced until later generations, such as S3 to S6. This
method requires less record keeping and works better where
the main objective is to improve quantitative traits such
as yield and earliness, rather than qualitative traits such
as flesh color and disease resistance. However, traditional
pedigree breeding is probably the more useful method for
watermelon since there are many qualitative traits that
can be selected in early generations. In that way, plants
or families having unsuitable traits that are simply inherited
(such as poor fruit flesh color) can be eliminated in early
generations. Otherwise, they would be carried along until
the S3 to S6 generation when field-testing would be practiced
in the single-seed-descent breeding method.
Backcross Breeding. Backcross breeding
is used to transfer one qualitative (highly-heritable) trait
into an otherwise superior inbred. The superior inbred is
referred to as the recurrent parent. Often, six generations
of selection and backcrossing to the recurrent parent are
used to recover the genotype of the recurrent parent (except
for the addition of the new trait) without the other undesirable
traits from the non-recurrent (donor) parent. Two versions
of the backcross method are used depending on whether the
gene of interest is recessive or dominant.
For the transfer of a trait controlled by a recessive gene,
the recurrent parent is crossed with the donor parent, and
the F1 backcrossed to the recurrent parent (Fig. 3.11).
In one scheme, the F1 is self-pollinated to produce the
F2, which will segregate for the trait of interest. Individuals
having the trait can then be backcrossed to the recurrent
parent to produce the BC1. The BC1 generation is then tested
for the trait, and individuals having it are self-pollinated
once again to produce a segregating generation for selection
and backcrossing to the recurrent parent. The process is
repeated until the BC6 generation when the best individuals
are self-pollinated and selected for the trait to produce
the improved inbred. The inbred does not need to be tested
extensively in trials, because it will be identical to the
original inbred, but with one new trait.
For the transfer of a trait controlled by a dominant gene,
the recurrent parent is crossed with the donor parent, and
the F1 backcrossed to the recurrent parent. The BC1 generation
is then tested for the trait, and individuals having it
are backcrossed to the recurrent parent. The process is
repeated until the BC6 generation when the best individuals
are self-pollinated and selected for homozygous expression
of the trait using progeny testing.
Inbred Development. The best selections
from the recurrent selection program should be self-pollinated
each cycle to begin inbred development. Pedigree selection,
and backcross breeding result in the production of elite
inbred lines. Each year, those inbred lines that are produced
from the different parts of the breeding program should
be increased by self-pollination, tested for useful horticultural
traits, and used in the production of tetraploid inbred
lines, as well as directly for the production of diploid
hybrids based on the traits they have, and what is needed
by the market.
Isolation blocks or screen cages can be used to make large
seed increases of the inbreds if that is needed. Isolation
blocks should be away from other watermelon fields, requiring
a separation of at least 1 mile. Bees should be provided
in the isolation block or cage by bringing in one strong
hive, unless there are sufficient numbers of wild bees.
Hybrid Testing. The final stage of breeding
is to produce hybrids for testing. Hybrids are usually made
between two monoecious inbreds. For triploid hybrid production,
the seed parent should have a distinctive rind pattern that
has recessive inheritance. For hybrid production with less
labor input, the seed parent could be male sterile. The
seed increase of the male sterile inbred would be accomplished
by pollinating male sterile plants with the heterozygote
(Ms ms) as the pollen parent. For seedless hybrid production,
the seed parent would be a tetraploid inbred.
Once they have been developed, all inbreds can be crossed
in all possible combinations. However, that might produce
too many entries to evaluate properly. For example, 20 inbreds
could produce (20 x 19)/2 = 190 different hybrids, without
including reciprocals. Thus, it may make more sense to make
hybrids only from pairs of inbreds having complementing
traits of the proper type.
Testing of experimental hybrids should progress in stages,
with fewer hybrids to test in later stages where more effort
is spent on each hybrid. The first year trials might have
two replications in each of two locations. In the second
year, the best hybrids could be evaluated in 8 to 12 locations
using the conditions available at each (grower fields, state
university experiment stations). In the third year, the
hybrids would be sent to grower trials throughout the production
regions of interest for trials involving 0.25 to 1.0 acre
using a total of 5-10 lb. of seeds for all trials. Seeds
should be screened for bacterial fruit blotch before sending
to growers. One can usually get good data from at least
10 of the 50 trials. Information from the 3 years of trialing
should lead to the release of the best one or two hybrids
in the fourth year.
Although there is not much advantage of hybrids over open-pollinated
varieties for most traits, it is thought that the former
are more uniform. Thus, it may be possible to get the same
yield in fewer harvests because of more uniform growth and
a more concentrated fruit set. Hybrids offer several advantages
over open-pollinated varieties. A major advantage is the
production of seedless triploids, which are produced by
crossing a tetraploid female inbred with a diploid male
inbred. Hybrids also can express heterosis, with the hybrid
performing slightly better than the best parent in some
cases. The amount of heterosis in watermelon is around 10%.
Another advantage is the ability to get an intermediate
fruit shape by crossing an elongate-fruited inbred with
a round-fruited one. Inbreds can be used to combine dominant
genes for resistance from each parent into a hybrid that
has more dominant genes expressed than either parent. A
hybrid that has large seeds for the grower to plant and
small seeds in the fruit sold to the consumer can be produced
by crossing a large-seeded female inbred with a small-seeded
male inbred. Finally, hybrids provide a way for the seed
company to protect their proprietary inbreds from theft.
The disadvantages of hybrids are that they add an extra
step to the breeding process, and increase the cost of seeds
since they are produced by hand pollination rather than
by bee pollination. Use of male sterile inbreds for seed
production should help reduce the cost of hybrid seeds in
the future.
Seedless Variety Development
Tetraploid Production. Use of triploid
hybrids has provided a method for production of seedless
fruit. The tetraploid method for seedless watermelon production
was invented by H. Kihara. He began development of tetraploids
in 1939, and had commercial triploid hybrids available 12
years later. The development of triploid varieties adds
several problems to the process of watermelon breeding:
extra time for the development of tetraploids; additional
selection against sterility and fruit abnormalities in tetraploid
lines; choice of parents for low incidence of hard seed
coats in the hybrids; the reduction in seed yield per acre;
reduced seed vigor for the grower; and the necessity for
the diploid pollenizer to use up to one-third of the grower's
production field.
Seedless varieties are produced by crossing a tetraploid
(2n=4x=44) inbred line as the female parent with a diploid
(2n=2x=22) inbred line as the male parent of the hybrid.
The reciprocal cross (diploid female parent) does not produce
seeds. The resulting hybrid is a triploid (2n=3x=33). Triploid
plants have three sets of chromosomes, and three sets cannot
be divided evenly during meiosis (the cell division process
that produces the gametes). This results in non-functional
female and male gametes although the flowers appear normal.
Since the triploid hybrid is female sterile, the fruit induced
by pollination tend to be seedless. Unfortunately, the triploid
has no viable pollen, so it is necessary to plant a diploid
variety in the production field to provide the pollen that
stimulates fruit to form. Usually, one third of the plants
in the field are diploid and two thirds are triploid, although
successful production has been observed with as little as
20% diploids. Varieties should be chosen that can be distinguished
easily so the seeded diploid fruit can be separated from
the seedless triploid fruit for harvesting and marketing.
Breeders interested in the production of seedless triploid
hybrids need to develop tetraploid inbred lines to be used
as the female parent in a cross with a diploid male parent.
One of the major limiting steps in breeding seedless watermelons
is the small number of tetraploid inbreds available. Development
of seedless hybrids will be discussed in the following stages:
(1) choice of diploid lines, (2) production of tetraploid
plants, (3) tetraploid line development, and (4) hybrid
production and testing.
Stage 1 involves choice of diploid lines to use in tetraploid
production. Most of the tetraploid lines being used by the
seed industry have gray rind so that, when crossed with
a diploid line with striped rind, it will be easy to separate
self-pollinated progeny (which will be seeded fruit from
the female parent line) from cross-pollinated progeny (which
will be seedless fruit from the triploid hybrid). The grower
should discard the gray fruit so they are not marketed as
seedless watermelons by mistake.
Stage 2 is the production of tetraploid plants. Many methods
have been used effectively in other crops to produce polyploids,
including tissue culture regeneration, temperature shock,
and X-rays. In watermelon, tetraploids can be produced routinely
using plants regenerated from tissue culture or using the
herbicide oryzalin. Colchicine (C22H25O6N) is a poisonous
alkaloid used in the treatment of gout. It is taken from the seeds and
bulbs of Colchicum autumnale and is a widely-used method for tetraploid production in
watermelon. Colchicine inhibits
spindle formation, and prevents separation of chromosomes
at anaphase. Of all the methods of colchicine application,
shoot apex treatment at the seedling stage was found most
effective.
For the seedling treatment method, the diploid line of
interest is planted in the greenhouse in flats (8x16 cells
is a popular size) on heating pads that keep the soil medium
at 85°F for rapid and uniform germination. When the
cotyledons first emerge from the soil, the growing point
is treated with colchicine to stop chromosome division and
produce a tetraploid shoot with four sets of chromosomes
rather than two. The colchicine solution is used at a concentration
of 1% for small-seed size varieties (‘Minilee’,
‘Mickylee’, ‘Sweet Princess’), 1.5-2.0%
for medium-seed size varieties (‘Allsweet’,
‘Crimson Sweet’, ‘Peacock Striped’,
‘Sugar Baby’), and 2-5% for large-seed size
varieties (‘Black Diamond’, ‘Charleston
Gray’, ‘Congo’, ‘Dixielee’,
‘Klondike Striped Blue Ribbon’, ‘Northern
Sweet’). Colchicine is applied to the seedling growing
point in the morning and evening for 3 consecutive days,
using 1 drop on small- or medium-seed size plants and 2
drops on large-seed size varieties. The treatment produces
plants that are diploid, tetraploid, or aneuploid, so it
is necessary to identify and select the tetraploids in later
stages. Treatment of the T0 diploids with colchicine results
in about 1% of the seedlings (referred to as T1 generation
tetraploids) being tetraploids. Some diploid varieties and
breeding lines produce a higher percentage of tetraploids
than others. For example, ‘Early Canada’ produces
many tetraploids and ‘Sweet Princess’ does not.
Tetraploids can be detected by the direct method of counting
chromosomes of cells under the microscope, or by comparing
stem, leaf, flower, and pollen size with diploid controls.
A popular method involves counting the number of chloroplasts
in stomatal guard cells using a leaf peel under the microscope.
Tetraploids have approximately 10-14 chloroplasts in each
guard cell (20-28 total on both sides of the stomate), whereas
diploids have only 5-6 in each guard cell (10-12 total).
The method is useful for screening many plants for ploidy
level in the seedling stage before transplanting to the
main part of the greenhouse or field nursery for self-pollination.
Usually, multiple methods are used, identifying tetraploid
seedlings using their phenotype in flats before transplanting,
the chloroplast number in the stomatal guard cells of the
true leaves in seedling flats and greenhouse pots, and by
the appearance of the fruit and seeds at harvest after self-pollination
in the greenhouse. Tetraploids usually have thicker leaves,
slower growth, and shorter stems than diploids.
Stage 3 involves tetraploid line development. Tetraploid
plants are selected (using methods such as leaf guard cell
chloroplast number) in the T0 generation (plants from colchicine
treated diploids) from the greenhouse flats where they were
treated with colchicine. It is then necessary to plant the
T1 generation in flats to verify that the plants are tetraploids
in that next generation, and transplant the selections to
greenhouse pots for self-pollination. Seeds from those selections
(T2) can then be increased in larger plantings such as field
isolation blocks to get sufficient numbers of seeds per
tetraploid line to use in triploid hybrid production.
The fertility and seed yield of tetraploid lines will increase
over generations of self- or sib-pollination, probably because
plants with chromosome anomalies are eliminated, resulting
in a tetraploid line with balanced chromosome number and
regular formation of 11 quadrivalents. Seed yield of tetraploid
lines in early generations is often only 50-100 seeds per
fruit and sometimes as low as 0-5 seeds compared to 200-800
seeds for diploids. Another problem with early generation
tetraploids is poor seed germination, making it difficult
to establish uniform field plantings. It may require as
much as 10 years of self-pollination before sufficient seeds
of tetraploid lines can be produced for commercial production
of triploid hybrids. Advanced generations of tetraploid
lines usually have improved fertility, seed yield, and germination
rate compared to the original lines. Some companies require
more than 100 lbs. of seed of a tetraploid inbred to be
available before beginning commercial production of the
triploid hybrid variety. Approximately 110 tetraploid plants
are required for production of each pound of triploid seeds.
Stage 4 is the evaluation of tetraploids (usually T3 generation
or later) as parents of triploid hybrids. The tetraploids
should be evaluated directly for rind pattern, high seed
yield, and other traits such as male sterility for reduced
hand labor in hybrid seed production. The major test for
tetraploids however, is as female parents in triploid hybrid
seed production after making controlled crosses using diploid
male parents. The resulting hybrids are tested in yield
trials with two rows of triploid plots alternating with
one row of diploid plots to assure adequate pollen for fruit
set in the triploid hybrids. Useful tetraploid inbreds should
produce triploid hybrids with excellent yield and quality
for the market type and production area of interest.
Triploid Evaluation. Evaluation of triploid
hybrids is similar to evaluation of diploid varieties already
discussed. There are a few special considerations, however.
Triploids are not inherently superior to diploids, so triploid
hybrids can be better or worse than their diploid parental
lines. Therefore, as in the case of diploid hybrids, many
combinations of parental lines should be evaluated in triploid
yield trials to identify the ones producing hybrids with
the best performance. In general, diploid inbred parents
that have poor horticultural performance will produce triploid
hybrids having poor performance.
One problem affecting triploid hybrids is empty seed coats
(colored or white) in the fruit. Under some environmental
conditions, fruit are produced with large obvious seed coats
that are objectionable to consumers. Triploid fruit should
be evaluated for seed coat problems during trialing. Some
selection should also be done on the parents before triploid
production. Seed coats will be large in the hybrids if the
parents have large seeds. Seed size is genetically controlled,
with at least three genes involved: l, s, and tss. Use of
tetraploid lines with small or tomato-size seeds may help
solve the problem. Besides genetic effects, certain unknown
environmental conditions seem to increase the number of
hard seed coats in poor performing triploid hybrids.
Commercial production of elite triploid hybrid seed is
done by hand in locations where labor is inexpensive, or
by bee pollination in isolation blocks. The tetraploid and
diploid inbreds are planted together in alternating rows,
or in alternating hills within each row. Where labor is
abundant, the staminate flowers can be collected from the
male (diploid) parent and used to pollinate the pistillate
flowers on the female (tetraploid) parent. Pollinated flowers
should be capped the previous day to keep bees out, then
covered after pollination to prevent self or sib-pollination
after the cross has been made. The flowers should be tagged
with the date so that the fruit can be harvested 35-50 days
later.
A method that requires less hand labor is to plant the
pollen and seed parents in alternating rows, and to remove
all staminate flowers from the seed parent rows during
flowering time, usually a period lasting several weeks.
Pistillate flowers on the female parent are tagged on the
day they open with the date to assure that the fruit are
mature when harvested, and to harvest only fruit that were
pollinated during the time staminate flowers were removed
from the female parent. Seeds that are harvested can also
be sorted mechanically for size, weight or density to separate
triploid seeds (resulting from cross pollination) from tetraploid
seeds (resulting from self- and sib-pollination).
When seed production is by bee pollination in isolation
blocks, the tetraploid flowers are sib- or cross-pollinated
84% of the time, producing 3x and 4x seeds (progeny). If
the 2x and 4x parents of the 3x hybrid have different rind
patterns, each of the three-ploidy levels can be distinguished
at harvest. For safety, the pollen parent plants should
be destroyed after fruit are set on the seed parent plants.
A useful combination is for the tetraploid parent to have
fruit with a gray rind pattern, and the diploid parent to
have fruit with wide stripes, so the resulting triploid
hybrid will have striped fruit, easily distinguished from
the gray fruited tetraploids that result from self- or sib-pollination
of the female parent.
Mechanization
The job of watermelon breeding can be made easier and more
efficient if mechanization is used for as many steps in
the process as possible. Small plot equipment can be used
for fieldwork to permit more germplasm to be tested with
fewer workers and at a lower cost. Small-plot seeders can
be used to plant seeds in the field with optimum seed spacing
and planting depth using fewer workers than if seeds are
planted by hand. If transplants are used to plant the test
plots, machine transplanters can be used to punch the hole
before the workers on the machine set the seedling into
the hole, and follow up with water and fertilizer after
the worker has pressed soil around the seedling, all while
riding down the field row. Seeds can be packeted using a
seed counter, and plot size can be optimized to gain the
maximum information for the lowest cost. Research indicates
that optimum plot shape is rectangular and block (replication)
shape is square. It is difficult to mechanize harvest since
it is done by hand, and each fruit is counted and weighed.
However, some efficiency can be gained by using portable
computers to collect and analyze data. In the advanced trials,
it is useful to estimate flesh sweetness (fruit soluble
solids content) using a refractometer, and rind toughness
using a spring-loaded punch or penetrometer.
If a greenhouse generation is used to expedite inbred development
or hybridization, automation systems are useful for handling
the many plants to be grown for self- or cross-pollination.
Such systems include automatic heating and cooling, drip
irrigation with fertilizer and/or other chemicals injected
into the water, trellis support for easy vertical training
of the plants, automatic overhead curtains to keep the greenhouse
from overheating during the day in the summer, and to keep
the greenhouse warmer at night in the winter. Computer systems
can provide efficient control of the greenhouse equipment
and help provide optimum conditions for plant growth.
For seed harvesting and handling, it is useful to have
a bulk seed extractor, washing screens, a seed sluice, and
seed dryers. Seed companies have used such machines for
years, and it is useful for the plant breeder to build smaller
versions that match the size of the plant breeding program. Watermelon breeding is a labor intensive job,
but mechanization can help make the most of the available
workers, funds, and time.