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Ipomoea

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DOI: 10.1007/978-3-642-21102-7_7

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Chapter 7
Ipomoea

Padma Nimmakayala, Gopinath Vajja, and Umesh K. Reddy

7.1 Introduction series. The cultivated sweetpotato is a hexaploid with

2n ¼ 90, Ipomea tiliacea and Ipomea tabascana are
tetraploids with ploidy level of 2n ¼ 60, and the rest
7.1.1 Origin and Distribution of the species are diploids containing 2n ¼ 30.
Sweetpotato was known to be originally domesti-
The genus Ipomoea (family Convolvulaceae) contains cated in the New World (Austin 1988; Zhang et al.
600–700 species including cultivated sweetpotato 2004) and further spread between latitude 42 N and
[Ipomea batatas (L.) Lam.]. Over half of them are 35 S, from sea level up to an altitude of 3,000 m. The
concentrated in the Americas, where there may be region between the Yucatán Peninsula of Mexico and
400 taxa, classified within the subgenera Eriosper- Orinoco River in Venezuela is postulated as the center
mum, Quamoclit, and Ipomoea. These three subgenera of diversity because of its rich morphological variation
contain ten sections; seven of them are mainly con- in cultivated sweetpotato collections as well as in four
fined to the northern and southern America, where other species in the Batatas group (Austin 1988).
they are distributed as cultigens, medicinal plants, Secondary centers of diversity are Peru and Ecuador.
and weeds (Austin and Huaman 1996). Many Ipomoea Archeological remains showed that the cultivated
species have considerable importance either as medic- sweetpotato existed in Peru and Mexico during
inal plants or as ornamental plants. Because of new 2500–2000 BC (Austin 1988). In addition, carbon-
discovery of more species and increased information dated (wild) specimens found in Peru were dated
on morphology, the series has been revised several back to 10000–8000 BC (Source: Natural History
times (Austin 1978, 1988, 1991; McDonald and Aus- Museum). The cultivated sweetpotato is known to be
tin 1990; Austin et al. 1993) and currently the series dispersed from the Americas in two waves. The ear-
Batatas contains, in addition to I. batatas, 13 wild liest migration was to Polynesia, although it is not
species classified as closely related to sweetpotato known when and how (Austin 1988). The second
(Table 7.1). All of these species except I. littoralis movement was by Columbus, who first introduced
are endemic to the Americas. Two are considered to sweetpotato to Europe during 1492, then to Africa in
be of hybrid origin. Ipomea leucantha has been deter- 1500. In the sixteenth century, Portuguese explorers
mined to be a cross derivative of I. cordatotriloba
introduced sweetpotato to Africa, India, southeastern
I. lacunosa hybrids, and I. grandifolia is known to be Asia, and the East Indies. Later on, Spanish trading
a hybrid derivative of I. cordatotriloba
I. batatas galleons introduced to Philippines (Austin 1988).
(Austin 1978). Cytological investigations have shown Studies on assessment of genetic diversity by using
that there are three different ploidy levels within the amplified fragment length polymorphism (AFLP)
markers traced pre-historic introduction to Oceania
from Mesoamerica (Rossel et al. 2000).
Nearly 26,000 accessions of Ipomoea spp. are
P. Nimmakayala (*)
available in 83 gene banks around the world (Rao
Gus R. Douglass Institute and Department of Biology, West
Virginia State University, Institute, WV 25112, USA et al. 1994). Out of these, about 8,000 accessions
e-mail: Padma@wvstateu.edu belong to the cultivated sweetpotato (Takagi 1988).

C. Kole (ed.), Wild Crop Relatives: Genomic and Breeding Resources, Industrial Crops, 123
DOI 10.1007/978-3-642-21102-7_7, # Springer-Verlag Berlin Heidelberg 2011
124 P. Nimmakayala et al.

Table 7.1 Ploidy, genome type, and self-compatibility of Ipomoea species from series Batatas
Ipomoea species 2n (x ¼ 15)a Genome typeb Self-compatibility
I. triloba 2x ¼ 30 A C
I. cordatotriloba (previously I. trichocarpa) 2x, 4x A C
I. cynanchifolia 2x ¼ 30 A C
I. lacunosa 2x ¼ 30 A C
I.
leucantha 2x ¼ 30 A C
I. umbraticola 2x ¼ 30 A C
I. ramosissima 2x ¼ 30 A C
I. tenuissima 2x ¼ 30 A C
I. grandifolia 2x ¼ 30 A C
I. littoralis 2x ¼ 30 B I
I. tiliacea 4x ¼ 60 X I
I. tabascana 4x ¼ 60 B C
I. trifida 2x, 3x, 4x, 6x B I
I. batatas 6x ¼ 90 B I
I. gracilis (outgroup) 4x ¼ 60 X I
C Self-compatible, I self-incompatible (Nishiyama et al. 1975)
a
Chromosome number from Jones (1974), Austin (1988) and Jarret et al. (1992)
b
Genome type from Jarret et al. (1992)

A comprehensive analysis of genetic variation and world production. The nutrient composition of the
distribution pattern is essential for sound conservation storage root starch is up to 33.3% (on fresh matter
strategies including sampling of extant genetic basis) and storage root sugar is as low as 0.6% (on
resources in germplasm collections, identification of fresh matter basis). Additionally, storage root contains
duplicates, selection for core collection, and future b-carotene, calcium, magnesium, iron, and zinc upto
explorations. About 200 accessions of sweetpotato 154, 4,091, 1,815, 10, and 6.3 ppm, respectively. This
are maintained in China consisting of Chinese land- corresponds to 15.4-mg b-carotene, 409-mg calcium,
races, new introductions, breeding lines, and cultivars 181-mg magnesium, 1-mg iron, and 0.6-mg zinc in
(Wang et al. 1998). The sweetpotato genebank held at 100 fresh storage roots (CIP 2009).
CIP, Peru, maintains a total of 5,526 cultivated acces-
sions from 57 countries, of which 2,589 are from Latin
America (Huamán and Zhang 1997).
7.2 Phylogenetic Relationships Among
Ipomoea Species in Batatas Section

7.1.2 Economic Importance In spite of importance, information pertaining to evo-

lution and phylogeny of cultivated sweetpotato is still
Sweetpotato is the seventh largest food crop with obscure. Several hypotheses have been put forward to
about 122 Mt produced annually worldwide (FAO- explain the origin of sweetpotato. Sweetpotato was
STAT 2006) and the fifth nutritional contributor to thought to be originated from the diploid I. leucantha
human diets in the developing countries (Plucknett Jacq., and further by polyploidization into a tetraploid
1991). It produces stable crop yields under a wide species, I. littoralis Blume (Nishiyama 1971). The
range of environmental conditions and one of the subsequent hybridization between these two
staple diets in many countries. Over 95% of the global species might have generated triploid Ipomeoa trifida
sweetpotato crop is produced in the developing (H.B.K.) Don., which finally settled to hexaploid by
countries, where it is the fifth most important crop on chromosome doubling. Another hypothesis for the
fresh weight basis after rice, wheat, maize, and cas- origin of sweetpotato may be through hybridization
sava. Asia is the world’s largest in sweetpotato pro- between I. trifida and I. triloba, resulting in wild
duction, with China alone producing 90% of total ancestor of I. batatas (Austin 1988). On the basis of
7 Ipomoea 125

morphology, ecology, cytology, and cluster analyses, in sweetpotato genotypic identification and classifica-
I. trifida and I. triloba were considered being the tion of genetic relationships.
closest extant relatives of the cultivated sweetpotato, The recent cpDNA data indicated that I. trifida is
which had supposedly arisen through an allopolyploid the most likely one of the diploid progenitors of hexa-
(Austin 1988). In a third hypothesis, the autopoly- ploid I. batatas (McDonald and Mabry 1992). On
ploidy of I. trifida was discussed as an alternative the basis of the available information on b-amylase,
pathway of origin (Shiotani 1987), and particularly a fairly conserved nuclear gene, Rajapakse et al.
the occurrence of 2n gametes (diplogametes) might (2004) cloned and sequenced b-amylase gene to
have been involved in the origin of the polyploidy study phylogenetic relationships on several species.
series, and allowed sexual interconnection among dif- In this study, I. tabascana, a tetraploid species with
ferent ploidy levels. In fact, the production of 2n B genome is sharing close relationship with I. trifida.
pollen was reported in diploid I. trifida and tetraploid I. tabascana, a species discovered by McDonald and
I. batatas (Orejda et al. 1990; Freyre et al. 1991). Austin (1990), was shown also to be closely related to
Therefore, a mechanism of sexual polyploidization sweetpotato. This study classified I. tiliacea as one of
through fertilization by non-reduced gametes is a the ancestral species, and the species I. triloba as not
strong possibility of gene flow among the species related to the cultivated sweetpotato as previously
with different ploidy levels. In summary, interspecific thought. Putting together various possible conclusions
hybridization involving the species, I. trifida (2x, 4x, based on investigations using genomic and organelle
6x), I. leucantha (2x), and I. littoralis (4x), and DNA, most closely related species of sweetpotato are
subsequent domestication gave rise to the cultivated I. trifida (2x) and I. tabascana (4x). In addition, fluo-
form of I. batatas (Nishiyama 1982). rescence in situ hybridization (FISH) data indicated
Many studies were performed using molecular that polyploidization was followed by decrease in the
genetic markers to resolve the phylogenetic relation- number of 18S rDNA loci in higher ploidy level and
ships of sweetpotato and its related species. The use of provided evidence for major genomic rearrangements
restriction fragment length polymorphism (RFLP, Jar- and/or diploidization of polyploid I. batatas. On the
ret et al. 1992), random amplified polymorphic DNA basis of chromosome morphology, tetraploid I. trifida
(RAPD, Jarret and Austin 1994), and microsatellites or appeared to be more closely related to sweetpotato
simple sequence repeat (SSR, Buteller et al. 1999; P. than I. tabascana (Srisuwan et al. 2006).
Nimmakayala et al. unpublished) markers revealed the
close relationship between I. trifida and I. batatas. A
total of 220 microsatellites (Nimmakayala et al.
Unpublished results) were used to understand the phy-
7.3 Cross-Compatibility Relationships
logenetic relationship of individual species (Fig. 7.1)
in the Batatas complex [I. batatas (6x), I. tabascana among Ipomoea Species
(4x), I. tiliacea (2x), I. trifida (4x), I. triloba (2x), and
I. leucantha (2x)]. The number of alleles as amplified The Ipomoea species in the Batatas complex were
by various species is presented in Fig. 7.2. The number divided into three subgroups (A, B, and X) based on
of alleles amplified in diploids, tetraploids, and hex- their selfing abilities, interspecific crossing capabil-
aploids was comparable based on the knowledge of ities as well as morphological and cytogenetic char-
their ploidy levels. This result indicated that the poly- acteristics (Ting et al. 1957; Jones 1965; Jones and
ploidization in Batatas complex followed massive Deonier 1965; Martin and Jones 1973; Nishiyama
genome reorganization and rearrangements through et al. 1975; Oracion et al. 1990). The A group
processes such as gaining or losing the alleles and includes I. triloba and three closely related species,
genomic fragments. Principal component analysis viz., I. lacunosa, I. trichocarpa, and I. ramoni. The
(PCA) was carried out and the results indicated dis- species within the A group are self-compatible and
tinct clustering pattern of various species (Fig. 7.3). cross-compatible with each other (Jones and Deonier
The construction of genetic relationships using 1965). The B group comprises the species I. batatas,
unweighted pair group method with arithmetic mean I. littoralis, and Ipomea trifida, and these species
(UPGMA) and PCA demonstrated further use of SSRs are self-incompatible but cross-compatible with each
126 P. Nimmakayala et al.

Fig. 7.1 Ipomoea species grown in the

green house (WVSU). (a) I. batatas a b
(6x); (b) I. tabascana (4x); (c) I. tiliacea
(2x); (d) I. trifida (4x); (e) I. triloba
(2x); (f) I. leucantha (2x)

c d

e f

Fig. 7.2 Number of alleles amplified using 220 I. Batatas specific SSRs in Batatas complex
7 Ipomoea 127

1
14 16
15
W1 Ipomoea setosa
W2 Ipomoea setosa
0.20 2 W3 Ipomoea tabascana
W4 Ipomoea tiliacea
W5 Ipomoea trifida
W6 Ipomoea trifida
5 W7 Ipomoea trifida
8 3
0.00 9 W8 Ipomoea trifida
6
W9 Ipomoea trifida
W10 Ipomoea triloba
W11 Ipomoea triloba
12 W12 Ipomoea triloba
0.20 4 10
W13 Ipomoea leucantha
1113 7 SP-2 Ipomoea batatas
SP-3 Ipomoea batatas
SP-4 Ipomoea batatas

0.40
–0.50 –0.17 0.15 0.48 0.80
Dim-1

Fig. 7.3 Principal component analysis of Batatas complex using microsatellite amplifications

other. The X group includes two well-known tetra- across the world mainly because of their high starch
ploids, I. tiliacea (4x) and I. gracilis (4x), which are content and resistance to two major nematodes.
self-incompatible but cross-compatible (Nishiyama Besides K123, many wild species related to sweetpo-
et al. 1975). The B group is cross-incompatible with tato have been introduced from Mexico, Guatemala,
the A and X group, but the A group is cross-compati- Colombia, Ecuador, and the United States. There are
ble with the X group when the latter is used as the many excellent disease- and pest-resistant traits in
pollen parent. Genome types of Ipomoea species as these species that can be used to improve sweetpotato
currently classified as presented in Table 7.1. Former (Iwanaga 1988; Komaki 2004; Zhang and Liu 2005).
Phylogenetic studies of the group based on morpho- Therefore, many researchers tried to make wide cross
logical traits (Austin 1988) indicated that cultivated to obtain interspecific hybrids by controlled pollina-
races of I. batatas (6x) share close relations with tions or somatic cell culture techniques.
I. trifida (HBK) G. Don (2x) and I. triloba L. (2x). However, interspecific hybrids between sweetpo-
However, the study of morphology in the series is tato and its related species have been scarce, mainly
difficult due to the overlapping of traits between spe- due to the genome differentiation and interspecific
cies, interspecific hybridization, and homoplasy. Aus- incompatibility (Martin 1970, 1982; Teramura 1979;
tin (1988) reported that, in the literature and in Shiotani et al. 1990; Lu and Li 1992). A few
germplasm collections, I. trifida is often confused B-genome species have been used to improve sweet-
with sweetpotato and that the two species are closely potato quality and disease resistance (Hozyo and Kato
related. Tetraploid samples previously identified as 1973; Iwanaga 1988; Iwanaga et al. 1991). Species in
I. trifida were later classified as I. batatas, based on the A-genome group may also be sources of genes for
flower morphology (Bohac et al. 1993). sweetpotato improvement. A-genome species such as
Utilization of wild germplasm in sweetpotato I. triloba, which can be found in both very dry and
breeding began in the year 1956 in Japan immediately extremely wet habitats (Martin and Jones 1973), may
after introduction of the wild hexaploid plants be a potential source of drought tolerance and resis-
(2n ¼ 90), designated K123 from Mexico (Nishiyama tance to root rots and foliar fungal diseases. So
and Teramura 1962; Nishiyama 1971). Subsequently, far, evaluation and interest in species belonging
K123-derived lines had attracted major attention to the A-genome group has been limited due to
128 P. Nimmakayala et al.

cross-incompatibilities, which have restricted their I. trifida (2n ¼ 4x ¼ 60): A tetraploid species, very
use in sweetpotato breeding (Wedderburn 1967). A- twining, spreading with dark purple leaves with the
genome species such as L. triloba, which can be found green tips. Leaves are cordite with sparse tip pubes-
in both very dry and extremely wet habitats (Martin cence. The flowering is limited (Fig. 7.1d).
and Jones 1973), may be a potential source of drought I. triloba (2n ¼ 2x ¼ 30): Spreading type with dark
tolerance and resistance to root rots and foliar fungal purple spots. The leaves are with sparse hairs and
diseases. Ting et al. (1957) performed more than 3,000 no lobes. The leaf petiole pigmentation was green-
interspecific crosses between the sweetpotato plant ish purple. The flowering is scarce (Fig. 7.1e).
and its 24 related wild species of the genus Ipomoea, I. leucantha (2n ¼ 2x ¼ 30): Mostly twining, spread-
yet not a single viable F1 hybrid was obtained. ing type with purple colored vines. The leaves are
Wedderburn (1967) made attempts at hybridization triangular in shape with no pubescence. The leaves
between I. batatas and I. trichocarpa or I. gracilis. were with slightly lobed with purple pigmentation
Only the initiation of embryo development occurred on abaxial surface. The flowering is very scarce
when I. trichocarpa as female parent was crossed with (Fig. 7.1f).
I. batatas as male parent, and the seeds produced by
this hybridization were not viable.

7.4.1 Confirmation of Ploidy Level

7.4 Description of Wild Species Ploidy level is determined directly by somatic chro-
of Sweetpotato mosome counts (Lower and Johnson 1969; Karp et al.
1982). Unfortunately, chromosome counts in sweet-
potato are difficult, due to the small size of chromo-
Wild species of sweetpotato are propagated by seeds.
somes and special difficulties associated with
No tuber formation has been found in the species of
chromosome staining and metaphase arrest (Jones
A and B groups of classification by Nishiyama et al.
et al. 1986). Ploidy level in sweetpotato, however,
(1975). The major problem in seed germination tests is
can also be indirectly determined by flow cytometry
the presence of the seed coat, which can delay or prevent
and counting chloroplasts in the guard cells. Among
imbibition, that is, hardseededness is a common prob-
the different methods, flow cytometry is the most
lem. The seeds are non-endospermic with axile foliar
powerful technique for measuring the DNA content.
embryos within hard seed coats (Steinbauer 1937).
It is an expensive, but quick and reliable tool to accu-
Consequently, scarification and chipping treatments
rately determine the ploidy levels. We determined
are useful in promoting germination. The scarification
ploidy levels of various species in Batatas complex
experiments with exposure of seeds to sulfuric acid for
using flow cytometry and counting chloroplasts in
30 min, followed by washing in running water, gave
guard cells (UK Reddy and coworkers personal com-
100% germination of seeds across different species. Six
munication).
species of Ipomoea are characterized as per the descrip-
tors for sweetpotato (Huamán 1991) as described below.
I. batatas (2n ¼ 4x ¼ 90): Cultivated sweetpotato 7.4.1.1 Chloroplast Count
with deep purple flowers (Fig. 7.1a).
I. tabascana (2n ¼ 4x ¼ 60): Greenish purple vines. Chloroplast number per guard cell pair was counted
The leaves are lobed with three deep lobes, sparse using the methodology developed by Compton et al.
pubescence and linear shaped. Flowers are pentag- (1996). Using fine forceps, the lower epidermis was
onal and purple colored with obovate and obtuse removed from fully expanded leaves of seedlings and
sepals. Flowering is very scarce (Fig. 7.1b). transferred to a microscopic slide in a drop of water
I. tiliacea (2n ¼ 2x ¼ 30): Spreading plant type with covered by a cover slip, and chloroplasts per guard cell
deep purple vines. The leaves are glabrous triangu- pair were counted under 400
magnification. Chlor-
lar in shape with no lobes with scarce flowering oplasts were counted in ten guard cell pairs per leaf
(Fig. 7.1c). and totally three different leaves per plant were used
7 Ipomoea 129

for counting. A total of five different diploid, tetra- Partec flow cytometer uses the DAPI (4, 6-diamidino-
ploid, and hexaploid accessions were included in the 2-phenylindole)-based staining solution, Cystain UV
study to minimize experimental error. The mean num- for staining the DNA of individual nuclei. The UV
ber and standard error of chloroplasts were calculated excitation of Cystain is provided by a mercury arc
for each accession in various species of the Batatas lamp and the blue emission of DAPI is collected by
complex (Fig. 7.4). Chloroplast counts across various filter. The Ploidy Analyzer displays the result as a
ploidy levels of different species of the Batatas com- DNA histogram at different channel representing dif-
plex indicated that the diploids had chloroplasts in the ferent ploidy levels in real time on a large LCD screen.
range of 9–12 per guard cell pair, and the tetraploid The machine was calibrated using a machine standard,
range was 14–22 per guard cell pair. This observation Partec DNA control UV (trout red blood cells) and the
is quite logical as the number of chloroplasts per guard nuclei isolated from leaf tissues of diploid species
cell pair in the tetraploids is expected to be more than served as an internal standard. The sample was
the diploids. In contrast, the expected chloroplast prepared following the standard protocol supplied by
number is approximately 2.5
in the hexaploids than the manufacturer (http://www.partec.de). The ploidy
the diploids. Intriguingly, in the current study, Nim- levels of various species in Batatas complex were
makayala and colleagues (unpublished) observed determined by flow cytometry. The results of various
chloroplasts per guard cell pair in the cultivated hexa- ploidy levels are presented in Fig. 7.5. The results
ploid species to be in the range of 10–14, which is obtained from five biological replications of diploids,
similar to the chloroplast number of diploid species. tetraploids, and hexaploids were analyzed. Interest-
ingly, the peak, which indicates the genome content,
is at the same position in all the samples of diploids
7.4.1.2 Flow cytometry and tetraploids (Fig. 7.5). In this experiment, when
various samples of hexaploids were analyzed, a peak
The ploidy level of various Ipomoea species was was noted at a different position than the diploids as
determined by flow cytometry using Ploidy Analyzer expected to be in the hexaploids. The position of the
PA I (Partec, Germany). The Ploidy Analyzer deter- tetraploid peak at the diploid position is not typical.
mines ploidy level by the precise measurement of total Out of the 13 species of Ipomoea section Batatas,
DNA content of the individual nuclei. It can measure I. trifida (HBK) Don (6x, K123), I. littoralis Blume
thousands of individual nuclei with a single pass. The (4x, K233), and I. leucantha Jacq (2x, K221) could

Fig. 7.4 Number of chloroplasts of ten guard cell pairs (Mean and standard error) across the Batatas complex
130 P. Nimmakayala et al.

Fig. 7.5 Histogram showing peaks generated by flow cytometry of diploid, tetraploid, and cultivated hexaploid sweetpotato

be crossed with sweetpotato (Nishiyama et al. combinations from I. triloba
I. trifida, and (I. tri-
1975; Shiotani and Kawase 1987); however, no pro- loba
I. lacunosa)
I. batatas (4x) by ovule culture
mising breeding lines could be selected. Kobayashi methods. The hybridization between sweetpotato
et al. (1994) obtained two interspecific hybridization and diploid I. trifida showed very low crossing rate
7 Ipomoea 131

(Komaki 2004). Somatic cell hybridization was also Compton ME, Gary DJ, Elmstrom GW (1996) Identification of
exploited to produce hybrids but only a few combina- tetraploid regenerants from cotyledons of diploid water-
melon cultured in vitro. Euphytica 87:165–172
tions were reported to be successful. For those hybrids FAOSTAT (2006) Food and Agriculture Organization of the
obtained, wild parents were among the following spe- United Nations, Production statistics. http://faostat.fao.org.
cies: I. triloba (Liu et al. 1994), I. lacunose (Zhang et al. Accessed 10 Sep 2010
2002), and I. cairica (Guo et al. 2006). Nevertheless, Freyre R, Iwananga M, Orejda G (1991) Use of Ipomoea trifida
(HBK.) G. Don germplasm for sweetpotato improvement. 2.
these somatic cell hybrid plants varied substantially in Fertility of synthetic hexaploids and triploids with 2n
their chromosome numbers, were morphologically gametes of I. trifida, and their interspecific crossability
leaned to their wild parent, and a few could set storage with sweetpotato. Genome 34:209–214
roots. By applying plant growth hormones to stimulate Guo JM, Liu QC, Zhai H et al (2006) Regeneration of
plants from Ipomoea cairica L. protoplasts and prodcuton
pollination, Cao et al. (2009) produced two novel F1 of somatic hybrids between I. cairica L. and sweetpotato,
interspecific hybrids between I. batatas (L.) Lam. I.batatas (L.) Lam. Plant Cell Tiss Org Cult 87(3):321–327
(2n ¼ 6x ¼ 90) and two wild species I. grandifolia Hozyo Y, Kato S (1973) The plant production of wild type
(2n ¼ 2x ¼ 30) and I. purpurea (2n ¼ 2x ¼ 30). plants in lpomoea trifida (H.B.K.) Don. Tokyo Natl Inst
Agric Sci Bull Ser D 24:35–60
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sweetpotato virus (SPV), while I. grandifolia is resis- Rome, Italy
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demonstrated that it is quite possible to integrate out- 29–38
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(H.B.K.) G. Don germplasm for sweetpotato improvement.
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