Cryptochromes

Adiantum capillus-veneris has five cryptochrome genes (Kanegae and Wada, 1998; Imaizumi et al., 2000) whereas Arabidopsis has only two (Li and Yang, 2006). Cryptochromes consist of a photolyase-related sequence in the N-terminus with a C-terminus extension. Trp-277, which is important for photolyase function, is not conserved in all fern cryptochromes, indicating that fern cryptochromes may not have any photolyase function, as is the case for Arabidopsis cryptochromes (Li and Yang, 2006). Even when fern cryptochromes were (1) expressed in photolyase-deficient E. coli, (2) exposed to UV light, and (3) irradiated with photoreactivating light (blue light for l hour), the blue light did not change the survival rate of the E. coli compared to the control without blue light irradiation. This result indicates that fern cryptochromes do not function as photolyase in E. coli cells (Imaizumi et al., 2000). The expression of each cryptochrome mRNA under a variety of light conditions and in different cell and gametophyte stages was studied (Imaizumi et al., 2000). For example, CRY3 mRNA expression level is higher in protonemata and sporophytes than in spores or prothallia, and CRY5 mRNA is mainly expressed in sporophytes. CRY4 is down regulated during spore germination by phytochrome. Intracellular localization was also examined precisely using a GUS-CRY construct introduced through particle bombardment. GUS-cry3 and 4 are clearly localized in the nucleus and GUS-cry4 is predominantly found in the nucleus. On the other hand, cryl, cry2, and cry5 are found in the cytoplasm (Imaizumi et al., 2000).

Intracellular photoreceptive sites of blue light responses were studied physiologically using a microbeam irradiator. These techniques showed that the blue light receptors mediating inhibition of spore germination (Furuya et al., l997) and promoting cell division (Wada and Furuya, 1978) should be localized in or very close to the nuclei. Ferns have two phototropins and one neochrome as blue light receptors, but these photoreceptors are mainly localized on the plasma membrane, not in the nucleus (Kadota et al., 1982; Sakamoto and Briggs, 2002; Kawai et al., 2003). Taken together, the blue light perception of these phenomena might be mediated by cry3 and/or cry4. However, we do not have any conclusive evidence for this hypothesis at the moment. To address this open question we need to screen for mutants deficient in blue light inhibition of redlight induced spore germination. We have tried to find spores which germinate under continuous blue light after the induction of germination by a red light pulse, but so far we have not succeeded. From phylogenetic analysis of the five CRY family genes, CRY 1 and CRY 2, and CRY 3 and CRY 4 consist of different subfamily groups, suggesting that CRY 3 and CRY 4 might function redundantly. That is a reasonable explanation for why there are no mutant spores defective in the blue light effect for germination.

1.11 Concluding remarks

Almost all the early developmental processes of fern gametophytes, such as cell growth, direction of cell growth, cell division and its timing, are controlled by light (branch formation (Wada et al., 1998) and negative phototropism of rhizoids (Tsuboi et al., 2006) have not been discussed in this chapter). Without light signals they are not able to progress to the next developmental stages, indicating that we can control early developmental stages synchronously with light. At early developmental stages, gametophytes consist of only two different cell types, a protonemal cell and a rhizoid, so they are very uniform. Since rhizoid cells do not differentiate further, gametophyte differentiation is restricted to protonemal cells. Because all the developmental processes mentioned above occur synchronously in protonemal cells, as far as we can see under a microscope, gametophytes are suitable for studying each of the processes of cell differentiation, at least from the stand points of cell biology, photobiol-ogy, and physiology. However, because molecular biological techniques are not yet established in fern gametophyte systems, few people are involved in fern studies. To advance fern studies, the top priorities are to (1) increase the number of people who study fern gametophytes, (2) establish molecular techniques in fern gametophytes, and (3) accumulate data such as EST for various stages of gametophyte development. A possible technique of gene introduction into spores through the spore coat during imbibition was reported using Marsilea (Klink and Wolniak, 2001) and Ceratopteris (Stout et al., 2003), but so far we have not replicated the technique using A. capillus-veneris. On the other hand, gene silencing using DNA fragments is available in A. capillus-veneris (Kawai-Toyooka et al., 2004) and Ceratopteris (Ratherford et al., 2004), similar to an RNA interference (RNAi) technique in various organisms. The technique is very simple and easy. PCR-amplified double stranded DNA fragments of a target gene (either cDNA or genome DNA) can be introduced into gametophyte cells with a hygromycin phosphotransferase gene driven by a cauliflower mosaic virus 35S promoter by particle bombardment. The effectiveness is different depending on the genes transformed but it is a relatively useful technique (Kawai-Toyooka et al., 2004). Expressed sequence tag (EST) libraries in ferns are available for A. capillus-veneris (Yamauchi et al., 2005) and Ceratopteris (Salmi et al., 2005). These techniques and databases of fern genomes are not yet used frequently, but may contribute to fern studies in the near future.

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