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Panopoulou, GD, Clark MD, Holland LZ, Lehrach H, Holland ND.  1998.  AmphiBMP2/4, an amphioxus bone morphogenetic protein closely related to Drosophila decapentaplegic and vertebrate BMP2 and BMP4: Insights into evolution of dorsoventral axis specification. Developmental Dynamics. 213:130-139.   10.1002/(sici)1097-0177(199809)213:1<130::aid-aja13>;2-z   AbstractWebsite

Amphioxus AmphiBMP2/4 appears to be a single gene closely related to vertebrate BMP2 and BMP4. In amphioxus embryos, the expression patterns of AmphiBMP2/4 suggest patterning roles in the ectodermal dorsoventral axis (comparable to dorsoventral axis establishment in the ectoderm by Drosophila decapentaplegic and vertebrate BMP4). In addition AmphiBMP2/4 may be involved in somite evagination, tail bud growth, pharyngeal differentiation (resulting in club-shaped gland morphogenesis), hindgut regionalization, differentiation of olfactory epithelium, patterning of the anterior central nervous system, and establishment of the heart primordium, One difference between the developmental role of amphioxus AmphiBMP2/4 and vertebrate BMP4 is that the former does not appear to be involved in the initial establishment of the dorsoventral polarity of the mesoderm, Dev. Dyn. 1998;213:130-139. (C) 1998 Wiley-Liss, Inc.

Holland, LZ, Kene M, Williams NA, Holland ND.  1997.  Sequence and embryonic expression of the amphioxus engrailed gene (AmphiEn): The metameric pattern of transcription resembles that of its segment-polarity homolog in Drosophila. Development. 124:1723-1732. AbstractWebsite

Vertebrate segmentation has been proposed as an evolutionary inheritance either from some metameric protostome or from a more closely related deuterostome, To address this question, we studied the developmental expression of AmphiEn, the engrailed gene of amphioxus, the closest living invertebrate relative of the vertebrates, In neurula embryos of amphioxus, AmphiEn is expressed along the anteroposterior axis as metameric stripes, each located in the posterior part of a nascent or newly formed segment, This pattern resembles the expression stripes of the segment-polarity gene engrailed, which has a key role in establishing and maintaining the metameres in embryos of Drosophila and other metameric protostomes, Later, amphioxus embryos express AmphiEn in non-metameric patterns - transiently in the embryonic ectoderm and dorsal nerve cord. Nerve cord expression occurs in a few cells approximately midway along the rostrocaudal axis and also in a conspicuous group of anterior cells in the cerebral vesicle at a level previously identified as corresponding to the vertebrate diencephalon. Compared to vertebrate engrailed expression at the midbrain/hindbrain boundary, AmphiEn expression in the cerebral vesicle is relatively late, Thus, it is uncertain whether the cerebral vesicle expression marks the rostral end of the amphioxus hindbrain; if it does, then amphioxus may have little or no homolog of the vertebrate midbrain, The segmental expression of AmphiEn in forming somites suggests that the functions of engrailed homologs in establishing and maintaining a metameric body plan may have arisen only once during animal evolution, If so, the protostomes and deuterostomes probably shared a common segmented ancestor.

Holland, PWH, Koschorz B, Holland LZ, Herrmann BG.  1995.  Conservation of Brachyury (T) genes in amphioxus and vertebrates: Developmental and evolutionary implications. Development. 121:4283-4291. AbstractWebsite

Homologues of the murine Brachyury (T) gene have been cloned from several vertebrates, and are implicated in mesoderm formation and in differentiation of the notochord, In contrast, the roles of the ascidian Brachyury gene may be restricted to presumptive notochord, To understand the evolution of Brachyury genes and their developmental roles, we have searched for homologues in amphioxus, representing the third chordate subphylum and the probable closest relative of the vertebrates. We report the isolation of two amphioxus cDNA clones with clear homology to Brachyury genes, and demonstrate that these derive from separate loci resultant from a recent gene duplication. This finding represents an exception to the emerging consensus of an archetypal prevertebrate genome in amphioxus, The spatial and temporal distribution of Brachyury transcripts during amphioxus development is remarkably similar to vertebrate Brachyury, in presumptive mesoderm, posterior mesoderm and the notochord, Gene expression extends throughout the anteroposterior axis of the notochord, despite the most rostral regions being a more recent specialization; it also persists into larval stages, despite differentiation into contractile tissue, We propose that roles of Brachyury in notochord differentiation are more ancient than roles in mesoderm formation, and that the latter are shared by cephalochordates and all vertebrates.

Holland, PWH, Garcia-Fernandez J, Holland LZ, Williams NA, Holland ND.  1994.  The Molecular Control of Spatial Patterning in Amphioxus. Journal of the Marine Biological Association of the United Kingdom. 74:49-60. AbstractWebsite

The embryology of amphioxus (Chordata: Cephalochordata) has features in common with vertebrate embryology, reflecting a dose phylogenetic relationship between the two taxa. Amphioxus differs from vertebrates, however, in having less complex organogenesis and cranial morphogenesis, and less specialization along the anteroposterior body axis. Here we illustrate this by describing the embryology of an amphioxus species, Branchiostoma floridae. To gain further insight into the origins, evolutionary divergence and comparative embryology of these taxa, we are comparing the molecular control of embryonic development in amphioxus and vertebrates. For these analyses, we are focusing on homeobox genes: a diverse multigene family implicated in developmental control in many Metazoa. We report the results of PCR-based experiments which reveal that the amphioxus genome has homeobox genes from several recognized gene classes. The PCR experiments also suggest that amphioxus has fewer 'Hox' and 'Msx' class homeobox genes than do vertebrates. We suggest, therefore, that amphioxus may be a living descendant from an intermediate stage in the evolution of homeobox gene family complexity, and the complexity of vertebrate developmental control. The pattern of gene expression during embryogenesis has been described for one amphioxus homeobox gene of the Hox class. This gene is primarily expressed in the presumptive neural tube of amphioxus neurulae, later embryos and larvae, in a spatially-restricted manner. The expression data lead us to suggest that Hox genes are involved in the control of spatial patterning in the neural tube of amphioxus; the data are also interpreted as giving insight into possible homology between the amphioxus and vertebrate body plans.

Holland, PWH, Holland LZ, Williams NA, Holland ND.  1992.  An Amphioxus Homeobox Gene - Sequence Conservation, Spatial Expression During Development and Insights into Vertebrate Evolution. Development. 116:653-&. AbstractWebsite

The embryology of amphioxus has much in common with vertebrate embryology, reflecting a close phylogenetic relationship between the two groups. Amphioxus embryology is simpler in several key respects, however, including a lack of pronounced craniofacial morphogenesis. To gain an insight into the molecular changes that accompanied the evolution of vertebrate embryology, and into the relationship between the amphioxus and vertebrate body plans, we have undertaken the first molecular level investigation of amphioxus embryonic development. We report the cloning, complete DNA sequence determination, sequence analysis and expression analysis of an amphioxus homeobox gene, AmphiHox3, evolutionarily homologous to the third-most 3' paralogous group of mammalian Hox genes. Sequence comparison to a mammalian homologue, mouse Hox-2.7 (HoxB3), reveals several stretches of amino acid conservation within the deduced protein sequences. Whole mount in situ hybridization reveals localized expression of AmphiHox3 in the posterior mesoderm (but not in the somites), and region-specific expression in the dorsal nerve cord, of amphioxus neurulae, later embryos and larvae. The anterior limit to expression in the nerve cord is at the level of the four/five somite boundary at the neurula stage, and stabilises to just anterior to the first nerve cord pigment spot to form. Comparison to the anterior expression boundary of mouse Hox-2.7 (HoxB3) and related genes suggests that the vertebrate brain is homologous to an extensive region of the amphioxus nerve cord that contains the cerebral vesicle (a region at the extreme rostral tip) and extends posterior to somite four. This proposed homology implies that the vertebrate brain probably did not evolve solely from the cerebral vesicle of an amphioxus-like ancestor, nor did it arise entirely de novo anterior to the cerebral vesicle.