Development of polarity and bilateral asymmetry in the palleal bud of Botryllus schlosseri (Pallas)

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Abstract
The development of the polarity and bilateral asymmetry of the future adult zooid has been traced to their earliest morphological expression in the palleal bud of Botryllus. The account is based upon continued observation of living buds. The polarized antero‐posterior and dorso‐ventral axes are first expressed by the skewing of a symmetrical, hemispherical bud towards the anterior end of the parental bud. Identification of these axes is reinforced by the development of a loop‐like blood circulation, the primary circulation, in the horizontal plane during the enlargement of the skewed hemisphere to form a stalked vesicle. Bilateral asymmetry is first expressed by the asymmetrical expansion of the vesicle stage. The right posterior corner of the vesicle expands further posteriorly and becomes more acute than the left posterior corner. This larger expansion persists throughout the development of the right atrial cavity, which finally expands across the mid‐line to partially surround the gut. The bilateral asymmetry, expressed in the expanded vesicle, is reinforced by the development in sequence of a rounded gut rudiment and a pericardial rudiment in positions that would be expected from the asymmetry of the expanded vesicle. The first appearance of the gut rudiment occurs earlier than had been recognized previously. Conflicting accounts of the time and mode of formation of the pericardial rudiment have been clarified. The results of this study are discussed in the context of determination of bud territory, polarity and bilateral asymmetry.