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into the possibility of transgenic approaches to
improve various traits. Drought tolerance has
been studied transgenically by insertion of the
yeast trehalose- 6- phosphate synthase 1 gene (Kon-
drák et al ., 2011), the Arabidopsis dehydroascor-
bate reductase gene (Eltayeb et al ., 2011), a betaine
aldehyde dehydrogenase ( BADH ) gene from
spinach (Zhang et al ., 2011), or constitutive ex-
pression of the potato StMYB1R- 1 transcription
factor (Shin et al ., 2011). In addition, Waterer
et  al . (2010) investigated the effect of wheat
mitochondrial Mn superoxide dismutase (SOD3:1),
barley dehydrin 4 (DHN 4), a canola transcrip-
tional factor ( DREB/CBF1 ), or a bromegrass
stress-inducible ROB5 gene in potato cultivar
Desiree. The transgenes were placed under the
control of either the constitutive P35S promoter
or a stress-inducible Arabidopsis COR78 pro-
moter. The promoter/transgene combinations,
COR78:DHN4 and COR78:ROB5, were the most
likely to enhance tuber yield under drought
stress. Kim et al . (2011) inserted into potato cul-
tivar Atlantic a peroxiredoxin gene ( 2- Cys Prx )
from Arabidopsis controlled by either the consti-
tutive P35S promoter or the stress-inducible
PSWPA2 promoter from sweet potato. The trans-
genics were more tolerant to heat shock than
untransformed controls, and the stress-inducible
promoter resulted in greater expression of the
transgene than P35S. Stacking antioxidant genes
may provide greater protection against multiple
stress conditions than deploying single genes
(Ahmad et al ., 2010). The strategy of using stress-
inducible promoters restricts expression of the
transgene to conditions when resistance is needed,
with the intention of economizing physiological
processes and avoiding the undesirable effects
of transgene expression during non-stress
conditions.
Improvement of salt stress tolerance of potato
has also been the target of considerable transgenic
research in recent years. Bayat et al . (2010) re-
ported improved in vitro salt tolerance for potato
cultivars transformed with a barley antiporter
gene, HvNHX2 . Overexpression of the GalUR gene,
an ascorbic acid pathway enzyme, provided potato
cultivar Taedong Valley with enhanced tolerance
to salt stress, as measured by growth and microtu-
berization of in vitro plantlets (Upadhyaya et al .,
2011). The Arabidopsis transcription factor, At-
DREB1A , under the control of a stress-inducible
Arabidopsis promoter ( rd294 ) in potato cultivar
Desiree, conferred salinity tolerance relative to
the level of expression of AtDREB1A under salt
stress conditions (Celebi-Toprak et al ., 2005;
Watanabe et al ., 2011). These initial studies of
transgenics using stress response genes, identi-
fied through the application of transcriptomic
and proteomic tools in model plants as well as in
potato, suggest a new wave of commercial trans-
genic potatoes with wide environmental adapta-
tion in the near future.
Resistance against biotic stress has been
emphasized in transgenic studies in recent years
as well. Defense against fungal pathogens has
been illustrated by the insertion of thionin genes
from brassicaceous species into potato, which
was reported to express increased resistance to
gray mold ( Botrytis cinerea ) (Hoshikawa et al .,
2012). The StoVe1 gene from an aubergine spe-
cies was overexpressed to reduce the infection
rate of Verticillium dahliae on potato cultivar De-
siree (Liu et al ., 2012). Likewise, expression of
both chitinase (chiA) and ribosome inactivating
protein (rip30) was combined to enhance the re-
sistance of potato to Rhizoctonia solani (M'Hamdi
et al ., 2012). In another study, the Rpi-bt1 resist-
ance gene from S. bulbocastanum under the con-
trol of a potato ubiquitin promoter conferred
resistance to late blight (Oosumi et al ., 2009).
A  new source of transgenic resistance to PVY
was found in the construction of a recombinant
heavy chain variable region (VH antibody) with
added hydrophobic residues targeting cytosolic
expression (Bouaziz et al ., 2009). Stacking of re-
sistance genes by inserting a double gene con-
struct ( Nicotiana tabacum AP24 osmotine and
Phyllomedusa sauvagii dermaseptin) into potato,
followed by retransformation with a Gallus gallus
lysozyme construct, yielded plants that were
highly resistant to Erwinia carotovora and in-
hibited fungal growth (Rivero et al ., 2012).
Pest resistance as well as pathogen resist-
ance has been the focus of several studies on
transgenic potato. The different crystal (cry) en-
dotoxins produced by the soil bacterium, Bacillus
thuringiensis , have been used to convey trans-
genic resistance to several insect pests of potato
(Pardo-López et al ., 2013). Various cry genes,
including cry1Ab under the control of a tuber-
specific, granule-bound starch synthase (GBSSi)
promoter (Kumar et al ., 2010), or a light-inducible
phosphoenolpyruvate carboxylae (PEPC) pro-
moter (Hagh et al ., 2009), and cryIIa1 with a
 
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