International Journal of Plant Breeding ©2007 Global Science Books

Breeding for Increased Sweetness in Sweet Corn

Kamol Lertrat1* • Taweesak Pulam2

1 Plant Breeding Research Center for Sustainable Agriculture, Faculty of Agriculture, Khon Kaen University, Khon Kaen, 40002, Thailand
2 Sweet Seed Co., Ltd., Prabuthabat, Saraburi, 18120, Thailand

Corresponding author: * [email protected]

ABSTRACT
Sweetness, a creamy texture and aroma, along with desirable kernel color, good germination, and high yield, are the important traits for
sweet corn. Various recessive mutants in corn expressed in the endosperm e.g. sugary1 (su1), sugary enhancer (se), shrunken2 (sh2),
brittle1 (bt) etc., which improve quality traits such as sweetness, flavor and nutritive value have been used either singly or in combination
for developing new commercial hybrids. In single mutant varieties, at harvest (20 days after pollination), the sugar (sucrose) concentration
in su1 and sh2 sweet corn is 3 and 8 times higher than wild type, respectively. For se-type when in combination with su1 the sucrose level
is as high as that in sh2. After harvest, the sugar in kernels of su1 and se types is rapidly converted to starch, but this conversion occurs
more slowly in sh2 type. However, both su1 and se sweet corn have more phytoglycogen or creamy texture, than sh2 variety without
difficulties in germination. The details of breeding methods using a combination of endosperm genes for sweetness improvement are
disclosed in the U.S. Patents Nos. 3,971,161 and 4,630,393. To date, two new high sugar types of commercial sweet corn, synergistic and
augmented sweet varieties have been developed and released by U.S. seed companies. Synergistic has a combination of su1, se, and sh2
kernels on each ear. It carries the seed quality and vigor of su1 varieties, the enhanced sweet corn flavor of se and harvest-ability and shelf
life approaching supersweet (sh2) type. Augmented sweet varieties, sh2 type, carry se modifier genes for tenderness and sweetness and the
sh2 gene for high sugars and long shelf life.
_____________________________________________________________________________________________________________

Keywords: augmented shrunken, eating quality, endosperm mutants, flavor improvement, quality improvement, supersweet corn,
synergistic, triple sweet
Abbreviations: WSP, water soluble polysaccharide

CONTENTS

INTRODUCTION........................................................................................................................................................................................ 27
ECONOMIC VALUE OF SWEET CORN .................................................................................................................................................. 27
TYPES OF IMPORTANT ENDOSPERM MUTANTS ............................................................................................................................... 28
THE DEVELOPMENT OF SWEET CORN BREEDING FOR SWEETNESS .......................................................................................... 28
Standard sweet corn varieties .................................................................................................................................................................. 28
Sugar-enhanced sweet corn varieties ....................................................................................................................................................... 28
Supersweet or extrasweet corn varieties .................................................................................................................................................. 28
High sugar sweet corn varieties ............................................................................................................................................................... 29
CONCLUSION ............................................................................................................................................................................................ 29
ACKNOWLEDGEMENTS ......................................................................................................................................................................... 29
REFERENCES............................................................................................................................................................................................. 29
_____________________________________________________________________________________________________________

INTRODUCTION corn (Evensen and Boyer 1986). It makes up most of flavor

and depends largely on kernel sucrose content. Texture is
Sweet corn (Zea mays L. var. rugosa Bonaf.) eaten in the determined primarily by pericarp tenderness, levels of water
immature stage, is widely used for human consumption soluble polysaccharides (phytoglycogen), and kernel mois-
throughout the world. It is an important source of fiber, ture content.
minerals, and certain vitamins. It is produced for three dis- The contents of this mini-review are about the economic
tinct markets; fresh, canning, and freezing. Production value of sweet corn, types of important endosperm mutants,
within these markets is largely independent of each other. and the development of sweet corn breeding for sweetness.
Newly developed products such as sweet corn milk and
soups are gaining popularity in many countries. New varie- ECONOMIC VALUE OF SWEET CORN
ties of sweet corn have been developed with improved con-
sistency, taste and shelf life. The adoption of newer “high According to the Food and Agriculture Organization of the
sugar or sweeter” varieties with longer shelf life and new United Nations statistic data, in 2005 the world’s sweet corn
sweet corn products have increased sweet corn consump- production for the fresh, frozen and canned food industries
tion and have helped to further expand the market. by harvested area is about 1.06 million ha. In 2004, the
Sweet corn eating quality of fresh or processed whole- world’s top four leading producers were the US (4.12 mil-
kernels, canned or frozen, is determined by its unique com- lion MT), Mexico (0.63 million MT), Nigeria (0.58 million
bination of flavor, texture, and aroma. Sweetness is the MT) and France (0.50 million MT). The world’s top four
most important factor in consumer satisfaction with sweet leading exporters of canned sweet corn are the US (0.14

Received: 28 February, 2007. Accepted: 9 April, 2007.

Invited Mini-Review
 International Journal of Plant Breeding 1(1), 27-30 ©2007 Global Science Books

million MT), Hungary (0.13 million MT), France (0.11 mil- pollination), contain 10.2% of sucrose and 22.8% of water
lion MT), and Thailand (0.10 million MT). And the world’s soluble polysaccharide (WSP), a creamy texture, about 3
top four leading exporters of frozen sweet corn are the US and 8 times the sugar and WSP contents of field corn, res-
(0.07 million MT), Hugary (0.04 million MT), New Zea- pectively (Creech 1965). The sugary varieties have a crea-
land (0.03 million MT), and Belgium (0.02 million MT). my texture and good corn flavor and are known for their
Global imports of canned sweet corn are valued over $468 good germination and seedling vigor. But their kernels can
million in 2004. The top five importing countries are the lose their sucrose from 14.4% to 5.7% (about 2.5 times) at
Russian Federation (0.09 million MT), Germany (0.06 mil- room temperature (27.0°C) 24 hours after harvest due to
lion MT), the UK (0.05 million MT), Japan (0.05 million sucrose rapidly converting to starch (Garwood et al. 1976).
MT), and the Republic of Korea (0.04 million MT). The These losses greatly affect the eating quality. As a result, the
global imports of frozen sweet corn were valued over $218 harvest and storage periods for the su1 varieties are short.
million. And, the top five importing countries for frozen These varieties are suitable for processing, e.g., canning and
sweet corn are the UK (0.07 million MT), Japan (0.05 mil- freezing.
lion MT), Belgium (0.02 million MT), the U.S. (0.02 mil- Most of sugary-type sweet corn breeding works for tem-
lion MT), and China (0.01 million MT). In terms of per ca- perate region have been done in the US. Dantings Early was
pita consumption sweet corn, the US currently consumes one of the first named sweet corn varieties that appeared in
2.5 kg per year higher than the EU (1.5 kg per year). And, 1844 (Galinat 1971) while Golden Bantam, released in 1902,
French per capita consumption is one of the highest in Eu- became one of the most important open-pollinated varieties.
rope (Hénard 2003). Many new hybrid varieties such as Jubilee, Silver Queen
(the most popular roadside market sweet corn variety in the
TYPES OF IMPORTANT ENDOSPERM MUTANTS US), Golden Cross Bantum, Early Sunglow, Merit, Seneca
Chief, Bonanza and Earlivee have been developed and mar-
Sweetness is affected by the amounts of sugar and starch in keted.
the endosperm. One of the areas of genetic improvement of
sweet corn has involved the selection of mutants that pro- Sugar-enhanced sweet corn varieties
duce high sugar levels in the kernel endosperm. To date,
starch biosynthesis and the genetic modification of endo- The se type varieties have a sugar content twice as much as
sperm carbohydrates have been researched and reviewed. normal sweet corn, extremely tender kernels, creamy texture
Several endosperm genes used in sweet corn improvement and good corn flavor. Sugar conversion occurs at the same
to increase sugar content and decrease starch content have rate as for the su1 type since the kernels contain more sugar,
been identified (Creech 1965; Boyer and Shannon 1984; their sweet taste retain longer after harvest. There are two
Tracy 1997). Four most useful mutants are shrunken2 (sh2), distinct groups within the varieties containing the se gene:
brittle1 (bt), sugary1 (su1), and sugary enhancer1 (se). homozygous and heterozygous. The homozygous varieties
The sh2 and bt located in chromosome 3 and 5, res- (sese su1su1) have higher sugar (20% to 35%) in 100% of
pectively are classified as class 1 mutants involved in large their kernels. While the heterozygous varieties (Sese su1su1)
reductions in starch and large increase in sugar. Mature dry have lower sugar (14% to 25%) and only 25% of the se type
kernels of both mutants are collapsed, angular, opaque and kernels and 75% of the su1 type therefore, homozygous
brittle. While the su1 and se located in chromosome 4 and 2, varieties are usually sweeter than heterozygous varieties
respectively are the class 2 mutants occurred later in the (Tracy 1997).
starch biosynthesis pathway and changed types and propor- High quality commercial se-hybrids have been deve-
tion of types of polysaccharides stored in the endosperm. loped and commercialized such as Seneca Arrowhead and
Mature dry kernels of the su1 are wrinkled and translucent. Temptation from Seminis Seeds; Sugar Buns and Miracle
While se’s kernels observed only in su1 lines, were inflated, from Crookham, Co.; and Spring Treat and Merlin from
light colored, slow drying, color varies with background, Mesa Maize, Inc.
collapsed, angular, opaque and brittle. The class 1 mutants However, progress in developing se hybrids has been re-
are epistatic to class 2 mutants (Boyer and Shannon 1984; latively slow due to the difficulty in identifying its presence
Tracy 1997). in dry seeds and its undesirable pale yellow kernel color.
Also, it appears that a number of recessive modifiers are re-
THE DEVELOPMENT OF SWEET CORN quired to attain high quality su1se hybrids (Tracy 1997).
BREEDING FOR SWEETNESS
Supersweet or extrasweet corn varieties
Standard or conventional sweet corn breeding method has
been used to develop commercial hybrids by breeders for The sh2 mutants at the immature milky stage (20 days after
more than century. This method involves the development pollination) contain 29.9% sucrose, the highest amount,
of homozygous inbred lines by selfing and selection of about 3 and 8 times the sugar content of the su1 (10.2%) and
desired phenotypes using pedigree selection from broad- field corn (3.5%), respectively. And it retains this level lon-
based breeding populations, the crossing of these inbred ger and has less starch than su1, but has a less creamy tex-
lines, and the evaluation of the hybrids from the crosses to ture due to very little WSP (Creech 1965). From Nelson’s
determine which have commercial potential. The most ef- study in 1980, at 21 days after pollination the bt kernels
ficient method used for flavor screening is bite taste at mar- contain 19.3% less sucrose than the sh2 kernels (22.5%) but
keting stage. higher than field corn (2.9%). The texture of supersweet
Sweet corn breeders have been extensively and success- corn is crispy rather than creamy as with the standard and
fully used 4 mutants, sh2, bt, su1, and se, singly or in combi- enhanced varieties. The fresh market shelf life is extended
nation in modern breeding to create new "high sugar" com- because of the slower conversion of sugars to starch after
mercial sweet corn varieties. The breeding work has been harvest. Seed kernels are lighter in weight, shrunken in ap-
centered on manipulation of these endosperm genes which pearance and difficult to germinate under cold soil tempe-
control the level of sugar found in the corn kernel. As a re- rature.
sults, 4 groups of sweet corn: standard, supersweet, sugary Supersweet corn sh2-type in the US was firstly deve-
enhanced, and high sugar sweet corn, have been developed loped by Professor John Laughnan, University of Illinois in
and successfully used for commercial production. the 1950’s. Illinois Foundation Seeds Inc. (IFSI) is the first
seed company to release a supersweet corn and it is called
Standard sweet corn varieties "Illini Xtra Sweet". In the early 1980s a widespread use of
the supersweet corn hybrid, from 2 to over 90% within five
Standard or traditional sweet corn varieties are su1 mutants. years, has started in the Florida growing area after a suc-
The su1 varieties, at immature milky stage (20 days after cessful marketing campaign by Abbott and Cobb Inc., to

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 Increasing sweetness in sweet corn: Breeding perspectives. Lertrat and Pulam

educate wholesale buyers, consumers and growers about have been released. These new types can be divided into
the superior qualities of supersweet corn. And the same two pollination groups as a supersweet or sugary variety
trend occurred throughout the US for all sweet corn grown (Tracy 2005).
for long-distance shipping. Simultaneously, the introduction Improved sh2 varieties (Augmented: Xtra-Tender
of new supersweet hybrids with necessary processing traits Brand®, Gourmet Sweet Brand®, and Multisweets®) have
marked the beginning of canned supersweet corn industry recently appeared. They have a sh2 endosperm that has been
(Pataky 2003). successfully combined with se genes and modifier genes.
Supersweet corn varieties, both sh2- and bt-types for the The kernels of these ears are more tender than those of ordi-
tropic were developed by Professor James Brewbaker, Uni- nary sh2 varieties. They are also very sweet (over 30% su-
versity of Hawaii in the 1960’s. Seventeen populations of gars), have a good corn flavor, good longevity and are more
sweet (su1 gene) and supersweet corn (bt, bt2 and sh2 genes) robust in the field than the sh2 varieties. Newly improved
have been bred in Hawaii for quality, yield, and tolerance to sh2 hybrids varieties have been developed and commercia-
maize mosaic virus and other diseases. These populations lized such as Xtra-Sweet and Xtra-Tender from Illinois
offer tropical breeders some of the best germplasm for Foundation Seeds Inc., Mirai® from Siegers Seed Co. and
sweet and supersweet corn improvement. Several of the va- Optimum and Holiday from Crookham, Co.
rieties are in commercial use internationally, as they have Synergistic varieties (Sweet BreedTM and Triple Sweet®)
been informally released during earlier cycles of breeding have been created by combining su1 gene with se and sh2
since 1968 (Thailand, sh2 COMP 1). All have a foundation genes. They are homozygous for the su1 gene but hetero-
in Hawaiian Sugar (tracing to Cuban × Golden Bantam), zygous for the other genes. The ear therefore contains dif-
with inputs from many different sources. The most widely ferent types of kernels.
grown OP populations are “Hawaiian Supersweet #9”, bt Sweet BreedTM such as Sweet Chorus, Sweet Rhythm,
and “Thai Supersweet”, a yellow sh2 population. Collabo- and Sweet Ice developed by Harris Moran Seeds are based
rative studies in Hawaii, Australia and Thailand have led to on a se heterozygous background have about 25% se kernels,
the development of commercially successful hybrids of 25% sh2 kernels and 50% su kernels. These varieties have
these sweet corns, which have expanding importance for good seed quality combined with higher sugar content than
fresh and processing markets in Asia. Widely grown super- the su1 varieties.
sweet hybrids include “Hawaiian Supersweet #10”, a 3-way Triple Sweet® varieties such as Avalon, Serendipity,
bt hybrid, Australia’s “H5”, a singlecross sh2 hybrid, and Honey select developed by Rogers seed, are based on a se
Thailand’s sh2 hybrids: ATS-5, Sugar-75, Hibrix3; bt: ATS- homozygous background have an ear composed of 75% su-
8. Several companies now market tropical supersweets, the gary enhancer kernels and 25% sh2 kernels. This type of
most extensive being that of Pulam’s “Sweet Seed Co.” in corn combines the outstanding tenderness and flavor of the
Thailand, and the products are well accepted by public se type with higher sugar content and an extended longevity
(Brewbaker et al. 2006). of the sh2 type. It also has good field robustness. Table-
These newer supersweet varieties are becoming the do- sweet® varieties such as Parfait TSW®, and Welcome TSW®
minant type in all major US sweet corn production regions developed by Mesa Maize, Inc. are the newly improved ho-
and many countries in Asia, Europe, South America and mozygous se varieties with high sugar contents and long
Africa. It is popularly known due to its higher sugar content shelf life.
and long shelf life, suitable for long-distance shipping. And
it has enabled manufacturers to can sweet corn without ad- CONCLUSION
ding sugar.
Because of the importance of sweet corn, breeders have
High sugar sweet corn varieties been devoted to the improvement of sweetness and others
valuable characteristics, such as yield, disease and insect re-
Although there are many commercial varieties of sweet sistance, and tolerance of environmental stress. By develop-
corn containing an individual gene of su1, se, sh2, or bt, ping new genetic combinations of four important endosperm
which provide a number of useful traits, these varieties are genes which are shrunken2 (sh2), brittle1 (bt), sugary1 (su1),
not without their problems. Sugary varieties lose their and sugary enhancer1 (se), new high sugar sweet corn vari-
sweetness rapidly after maturity. Supersweet (sh2, or bt) eties of great texture and taste with long shelf life have been
varieties have a tough pericarp, lack creaminess, poor corn developed and marketed. These high sugar sweet corn with
flavor and poor germination. Sugary enhanced varieties still extending postharvest quality hybrid varieties are well ac-
have lost their sweetness faster than in supersweet varieties. cepted by consumers both in the US and Japan. The world-
For this reason, breeders have developed new varieties with wide expansion of using the high sugar sweet corn hybrids
a high sugar content, good germination and superior agro- especially for fresh market can be anticipated. However,
nomic traits by using genetic combinations of other endo- better varieties with an emphasis on superior kernel quality
sperm genes with the sugary allele. Details of the breeding and agronomic requirements such as ear size and shape, ker-
methods are disclosed in the U.S. Patents Nos. 3,971,161 nel color and size unique to each specific market is required.
and 4,630,393. The sweet corn breeding program must go on and need
Bonucci, US Patent Nos. 3,971,161 and 4,630,393 has more support.
developed two new sweet corn hybrids, “Sweet Gene hyb-
rid” and “Sweetie”. The first hybrid or the sugary-shrunken ACKNOWLEDGEMENTS
hybrid, having the genotype su1su1Sh2sh2 is derived from a
cross between sugary-shrunken inbred (su1su1 sh2sh2) as fe- We would like to sincerely thank Professor James Brewbaker from
male with sweet corn male inbred (su1su1Sh2Sh2). The F1 is the University of Hawaii in the United States for laying the foun-
normal appearing sweet corn seed and produced F2 ears dation for sweet corn breeding in Thailand and for providing the
characterized by a 3:1 segregation of sugary (su1su1) ker- valuable information used in this review.
nels to shrunken (sh2sh2) kernels. The hybrid has very high
in both sugars (approximately 50% more sucrose, 33%
REFERENCES
more total sugars) and WSP. The second hybrid, “Sweetie”
or “Improved Supersweet” hybrid, having the genotype
Bonucci PA (1986) Production of hybrid “improved supersweet” sweet corn.
Su1su1 sh2sh2 is derived from a cross between sweet corn US Patent Number: 4,630,393
inbred (Su1Su1 sh2sh2) as female with sugary-shrunken male Bonucci PA (1979) Production of hybrid sweet corn. US Patent Number:
inbred (su1su1 sh2sh2). 3,971,161
After breeders have been “stacking” the endosperm Brewbaker JL (2006) Development of supersweet maize adapted to the tropics.
genes, new varieties contain a number of different combi- In: Mercer CF (Ed) Breeding for Success: Diversity in Action, Australasian
nations of the three major genes and their modifier genes Plant Breeding Conference Christchurch, New Zealand, 18-21 April 2006, pp

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 International Journal of Plant Breeding 1(1), 27-30 ©2007 Global Science Books

44-49 Hénard MC (2003) France processed sweet corn annual 2003. USDA Foreign
Boyer CD, Shannon JC (1984) The use of endosperm genes for sweet corn Agricultural Service, Global Agriculture Information Network Report Num-
improvement. In: Janick J (Ed) Plant Breeding Reviews (Vol 1), John Wiley ber: FR3054, 5 pp
and Sons, Inc, pp 193-161 Nelson OE (1980) Genetic control of polysaccharides and storage protein syn-
Creech RG (1965) Genetic control of carbohydrate synthesis in maize. Gene- thesis in the endosperm of barley, maize, and sorghum. Advances in Cereal
tics 52, 1175-1186 Science and Technology 3, 41-71
Evensen KB, Boyer CD (1986) Carbohydrate composition and sensory quality Pataky JK (2003) Fifty years of super supersweet corn: It keeps getting better.
of fresh and stored sweet corn. Journal of the American Society for Horticul- Available online: http://agronomyday.cropsci.uiuc.edu/2003/supersweet_corn/
tural Science 111, 734-738 Tracy WF (1997) History, genetics, and breeding of supersweet (shrunken2)
Galinat WC (1971) The evolution of sweet corn. Bulletin no. 591. University sweet corn. In: Janick J (Ed) Plant Breeding Reviews (Vol 14), John Wiley
of Massachusetts Agricultural Experiment Station, 20 pp and Sons, Inc, pp 189-236
Garwood DL, McArdle FJ, Vanderslice SF, Shannon, JC (1976) Postharvest Tracy B (2005) The cornfusion about sweet corn. University of Wisconsin.
carbohydrate transformations and processed quality of high sugar maize ge- Available online: http://www.uwex.edu/ces/sweetcorn/UWsweetpages/Corn
notypes. Journal of the American Society for Horticultural Science 101, 400- fusion.htm
404

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