The 70-kD heat shock proteins (Hsp70s) are highly conserved molecular chaperones that play essential roles in cellular processes including abiotic stress responses. Physcomitrella patens serves as a representative of the first terrestrial plants and can recover from serious dehydration. To assess the possible relationship between P. patens Hsp70s and dehydration tolerance, we analyzed the P. patens genome and found at least 21 genes encoding Hsp70s. Gene structure and motif composition were relatively conserved in each subfamily. The intron-exon structure of PpcpHsp70-2 was different from that of other PpcpHsp70s; this gene exhibits several forms of intron retention, indicating that introns may play important roles in regulating gene expression. We observed expansion of Hsp70s in P. patens, which may reflect adaptations related to development and dehydration tolerance, and results mainly from tandem and segmental duplications. Expression profiles of rice, Arabidopsis and P. patens Hsp70 genes revealed that more than half of the Hsp70 genes were responsive to ABA, salt and drought. The presence of overrepresented cis-elements (DOFCOREZM and GCCCORE) among stress-responsive Hsp70s suggests that they share a common regulatory pathway. Moss plants overexpressing PpcpHsp70-2 showed salt and dehydration tolerance, further supporting a role in adaptation to land. This work highlights directions for future functional analyses of Hsp70s.
MYB transcription factors (TFs) are one of the largest TF families in plants to regulate numerous biological processes. However, our knowledge of the MYB family in Physcomitrella patens is limited. We identified 116 MYB genes in the P. patens genome, which were classified into the R2R3-MYB, R1R2R3-MYB, 4R-MYB, and MYB-related subfamilies. Most R2R3 genes contain 3 exons and 2 introns, whereas R1R2R3 MYB genes contain 10 exons and 9 introns. N3R-MYB (novel 3RMYB) and NR-MYBs (novel RMYBs) with complicated gene structures appear to be novel MYB proteins. In addition, we found that the diversity of the MYB domain was mainly contributed by domain shuffling and gene duplication. RNA-seq analysis suggested that MYBs exhibited differential expression to heat and might play important roles in heat stress responses, whereas CCA1-like MYB genes might confer greater flexibility to the circadian clock. Some R2R3-MYB and CCA1-like MYB genes are preferentially expressed in the archegonium and during the transition from the chloronema to caulonema stage, suggesting their roles in development. Compared with that of algae, the numbers of MYBs have significantly increased, thus our study lays the foundation for further exploring the potential roles of MYBs in the transition from aquatic to terrestrial environments.
Senescence-associated receptor-like kinase (SARK) family members in Arabidopsis, soybean, and rice are known to be positive regulators of leaf senescence. In the meantime, SARKs are extensively involved in stress response. However, their function and underlying molecular mechanism in stress responses in moss are not well known. Here, we investigated functional roles of SARK isolated from Physcomitrella patens (PpSARK) in salt stress response and senescence. PpSARK transcripts significantly accumulated under NaCl and abscisic acid (ABA) treatments, with higher expression in the moss gametophyte stage. Insertional gain-of-function mutants of PpSARK (PpSARKg) were more tolerant to salt stress and ABA than wild type (WT), whereas senescence of mutants was delayed during the protonema stage. Expression of stress-responsive genes in the ABA related pathway, such as PpABI3, PpABI5, PpPP2C, and PpLEA were significantly higher in PpSARKg and WT under salt stress conditions, suggesting that PpSARK might positively regulate salt tolerance via an ABA-related pathway. Endogenous ABA contents also increased 3-fold under salt stress conditions. These results indicate that PpSARK functions as a positive regulator in salt stress responses, while possibly functioning as a negative regulator in senescence in moss.
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