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Rapid evolutionary divergence of Photosystem I core subunits PsaA ...

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136<br />

Figure 2. Phylogenetic analysis <strong>of</strong> concatenated <strong>PsaA</strong> and PsaB proteins using the neighbor-joining distance method. For this analysis, all<br />

gaps created by the alignment were excluded. The first number at nodes represent the bootstrap percentage from 100 replicates for the<br />

neighbor-joining distance method, the second number indicates the bootstrap support for this node found by the maximum parsimony method.<br />

Values below 50% are indicated by – or not shown. Synechocystis PCC6803 was arbitrarily chosen as an outgroup. The scale bar indicates<br />

substitutions per amino acid position.<br />

The neighbor-joining <strong>PsaA</strong>/B tree exhibits unexpectedly<br />

long branches within the Synechocococcus/Prochlorococcus<br />

cluster (Figure 2), despite the<br />

fact that these species are phylogenetically close. This<br />

suggests considerable differences in the <strong>evolutionary</strong><br />

rates <strong>of</strong> the psaA/B operon both in between marine<br />

cyanobacteria and between these and the other phototrophs.<br />

The high <strong>divergence</strong> in PS I protein sequences<br />

between these three strains is confirmed by the comparison<br />

<strong>of</strong> sequence identities (Table 1). The <strong>PsaA</strong> sequences<br />

are more divergent than the PsaB sequences.<br />

Prochlorococcus sp. SS120 shows the highest deviation<br />

<strong>of</strong> both the <strong>PsaA</strong> and PsaB sequences. For<br />

both <strong>PsaA</strong> and PsaB, Heterocapsa showed by far the<br />

highest deviation (not shown). Since the larger number<br />

<strong>of</strong> insertions and deletions in these proteins from Heterocapsa<br />

complicated the alignment, leading to more<br />

gaps, they were excluded from the Table. Compared<br />

to the species indicated in Table 1, <strong>PsaA</strong> from H. triquetra<br />

showed sequence identities between 51.4% to<br />

P. marinus SS120 and 56.6% to Cyanophora paradoxa.<br />

For Heterocapsa PsaB, the sequence identity<br />

ranged between 51.3% to Prochlorococcus sp. MED4<br />

and 55.9% to Cyanophora paradoxa.<br />

Discussion<br />

The amino acid sequence analysis <strong>of</strong> the PS I <strong>subunits</strong><br />

<strong>PsaA</strong> and PsaB from one Synechococcus strain<br />

(WH 7803), representative <strong>of</strong> the Marine-Cluster A<br />

(Waterbury and Rippka 1989), and two Prochlorococcus<br />

strains, representative <strong>of</strong> the low-light (SS120)<br />

and high-light adapted (MED4) ecotypes (Moore et al.<br />

1995, 1998; Urbach et al. 1998), are in agreement with<br />

previous molecular studies showing that these marine<br />

oxyphototrophs are closely related and probably<br />

evolved from a common ancestor (Hess et al. 1995;<br />

Urbach et al. 1998; Honda et al. 1999; Turner et al.<br />

1999).<br />

The high <strong>evolutionary</strong> rate <strong>of</strong> the <strong>PsaA</strong> and PsaB<br />

sequences from all the three marine oxyphototrophs

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