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Fig. 17.2 Models of multiple interactions between ABA and other plant hormones in cold
stress response. There are multiple points of interaction between ABA and other plant hormones
including GA ( a ), cytokinin ( b ), ethylene ( c ), JA ( d ), and SA ( e ) in regulation of plant response
to cold stress. CTK cytokinin, ETH ethylene, DELLA DELLA protein, AHK Arabidopsis histi-
dine kinase, AHP Arabidopsis histidine phosphor-transferase, ARR Arabidopsis response regu-
lator, EIN3 Ethylene insensitive3, JAZ jasmonate-zim-domain protein, MYC2 MYC-related
transcriptional activator2
responses in both ABA-dependent and ABA-independent signalling pathways
(Tran et al. 2007 ); on the other hand, the homologous gene AHK1 was identi-
fied as an osmotic stress sensor that positively regulates abiotic stress (Wohlbach
et al. 2008 ; Kumar et al. 2013 ; Tran et al. 2007 ). Increasing evidence suggests a
negative role for cytokinin signalling in cold- and ABA-mediated abiotic stress.
For instance, ahk2 ahk3 double mutants are highly tolerant to cold, drought, and
salt stress and show strong expression of ABA-responsive genes (Jeon et al. 2010 ;
Tran et al. 2007 ). However, whether cytokinin receptors can perceive stress signals
is still unclear. AHP2, AHP3, and AHP5 function as redundant negative regula-
tors of the drought stress response in both ABA-dependent and ABA-independent
manners (Nishiyama et al. 2013 ). These studies support the notion that AHK2-
AHK4 and AHPs play negative roles under unfavourable environmental conditions
and may integrate with ABA signalling to modulate stress responses (Fig. 17.2 b).
Type-A ARR genes are negative response regulators in cytokinin signalling path-
way. They are induced by cytokinin as well as by cold stress (To et al. 2004 ).
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