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An amphioxus LIM-homeobox gene, AmphiLim1/5, expressed early in the invaginating organizer region and later in differentiating cells of the kidney and central nervous system.
Langeland JA
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Holland LZ
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Chastain RA
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Holland ND
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A LIM-homeobox gene, AmphiLim1/5, from the Florida amphioxus (Branchiostoma floridae) encodes a protein that phylogenetic analysis positions at the base of a clade comprising vertebrate Lim1 and Lim5. Amphioxus AmphiLim1/5 is expressed in domains that are a composite of those of vertebrate Lim1 and Lim5, which evidently underwent subfunctionalization after duplication of an ancestral protochordate Lim1/5. During amphioxus development, transcription is first detected in the ectoderm of the blastula. Then, in the gastrula, a second expression domain appears in the mesendoderm just within the dorsal lip of the blastopore, a region known to have organizer properties in amphioxus. This mesendodermal expression corresponds to Lim1 expression in the Spemann organizer of vertebrates. At least one of the functions of vertebrate Lim1 in the organizer is to control the transcription of genes involved in cell and tissue movements during gastrulation, and a comparable early function seems likely for AmphiLim1/5 during gastrular invagination of amphioxus. Later embryos and larvae of amphioxus express AmphiLim1/5 in clusters of cells, probably motoneurons, in the anterior part of the central nervous system, in the hindgut, in Hatschek's right diverticulum (a rudiment of the rostral coelom), and in the wall of the first somite on the left side (a precursor of Hatschek's nephridium). In the early larva, expression continues in neural cells, in Hatschek's nephridium, in the wall of the rostral coelom, in the epidermis of the upper lip, and in mesoderm cells near the opening of the second gill slit. The developmental expression in Hatschek's nephridium is especially interesting because it helps support the homology between this amphioxus organ and the vertebrate pronephros.
Figure 1. Deduced amino acid sequence of AmphiLim1/5 from the Florida amphioxus, Branchiostoma floridae, with two LIM domains (each boxed), a homeodomain (single underlined), an arginine-rich region (double underlined), and a tyrosine-rich region (bracketed). The arrowheads correspond to intron positions in the base sequence.
Figure 2. Neighbor-Joining phylogenetic analysis of amphioxus AmphiLim1/5 protein in the context of closely related LIM-homeodomain proteins from other chordates. Tree topology with bootstrap support based on 1000 replicates is constructed with Clustal_X.
Figure 3. AmphiLim1/5 expression in developing amphioxus. Whole mount side views with anterior toward left. Sections are viewed from posterior end of animal. Whole mount and section scale lines respectively 50 μm and 25 μm. A) Blastula with expression in animal hemisphere. B) Mid-gastrula in blastopore view with expression in dorsal quadrant. C) Section in plane of dashed line in B; expression in ectoderm and dorsal mesendoderm (arrow). D) Side view of late gastrula with expression in neural plate and in posterodorsal mesendoderm. E) Section through a in D showing expression in neural plate. F) Section through b in D showing expression in dorsal mesendoderm. G) Side view of mid-neurula with expression in central nervous system, hindgut, rudiment of Hatschek's right diverticulum (asterisk), and first somite on left side (arrow). H) Section through a in G showing expression in anterior neural plate, epidermis on left side, Hatschek's right diverticulum (single arrow), but not in Hatschek's left diverticulum (twin arrow). I) Section through b in G showing expression on epidermis on left side and in most anterior somite on left side (arrow). J) Section through c in G showing expression in hindgut, but not in notochord (arrow). K) Dorsal view of mid-neurula showing expression in central nervous system, hindgut, and first somite on left side (arrow). L) Side view of early larva with expression in cells of dorsal nerve cord, rostral coelom (single arrow), Hatschek's nephridium (twin arrows), mesoderm associated with rudiment of second gill slit (asterisk), and neurenteric canal (arrowhead) of tail bud. M) Section through a in L showing expression in ventral and lateral cells of central nervous system and in rostral coelom (arrow). N) Section through b in L showing expression in ventral and lateral cells of central nervous system, in epidermis on left side, but not in Hatschek's left diverticulum (arrow). O) Section through c in L showing expression in epidermis on left side and in wall of Hatschek's nephridium (arrow). P) Section through d in L with expression in mesoderm cells near rudiment of second gill slit. Q) Side view of anterior end of later larva with expression in Hatschek's nephridium (single arrow) and in epidermis bordering upper lip of mouth (twin arrows). R) Section through a in Q, at level of mouth (arrow), showing expression in upper lip epidermis and in underlying Hatschek's nephridium.
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