The growth of supernumerary legs in the cockroach.

When the anteroposterior axis of a cockroach leg is reversed at a graft by exchanging a left leg for a right leg at the mid-tibia level, regeneration occurs in the region of the graft/host junction. This results in the formation of a pair of lateral supernumerary legs. In these experiments the patterns of cell division which take place during supernumerary leg formation were observed in sections of regenerating legs of the cockroach Leucophaea maderae. Early patterns of cell division resemble those seen in control grafts in which no axial reversal had been carried out during grafting. These cell divisions are associated with the process of wound healing. Later, a large area of the epidermis proximal to the graft/host junction becomes activated and shows a rapid rate of cell division. This area forms two outgrowths which grow by cell division throughout their epidermis to form the epidermis of the supernumerary legs. The results are more consistent with the view that the formation of supernumerary legs involves dedifferentiation of the epidermis in the region of the graft/host junction to form a blastema, rather than being due to local cell division at the point of maximum pattern discontinuity. This conclusion is used to offer an explanation for the range of different types of outcome of left-right grafts that has been observed.

Separation of wound healing from regeneration in the cockroach leg.

It has been shown that after a critical point in the moult cycle of a cockroach, wound healing can occur but regeneration of pattern does not take place until the following intermoult period. Leg removal after the critical point is used to separate the processes of wound healing and leg regeneration. This permits the study of patterns of cell division resulting from wound healing to be distinguished from those involved in leg regeneration. During wound healing, cell division occurs in the epidermal cells of approximately the distal half of the trochanter. The cells then return to the resting state until after the next ecdysis. Regeneration starts with cell division occurring in the distal half of the trochanter, and then spreading to include cells of the proximal trochanter and distal coxa. This spread and the following patterns of growth and redifferentiation appear to be the same as for regeneration following leg removal prior to the critical point, with the more distal structures completing early stages of regeneration first. Scanning electron micrographs of the cuticle of the trochanter after the ecdysis following leg removal support the evidence from the patterns of cell division in suggesting that the distal half of the trochanter is dedifferentiated during wound healing.

Blastema formation and cell division during cockroach limb regeneration.

Recent models of pattern formation in insects have been derived largely from observations on regenerated cuticular patterns. Such models make assumptions about the behaviour of the underlying epidermal cells, their movement and patterns of cell division. The present study, designed to test these assumptions, looks at the patterns of wound healing and cell division after amputation at the trochanter-femur joint of the metathoracic leg in the cockroach. It shows that the wound is closed by cell migration and that regeneration occurs by dedifferentiation of the trochanter and distal coxa to form a blastema which grows and redifferentiates to form the new limb. The extent of the spread of dedifferentiation is confirmed by a scanning electron microscope study of the coxa after the moult following amputation. The results highlight the need for a greater knowledge of cell behaviour during pattern formation before we can begin to understand the processes involved in pattern formation.

Naturally occurring abnormalities (Bruchdreifachbildungen) in the chelae of three species of Crustacea (Decapoda) and a possible explanation.

Naturally occurring abnormalities (Bruchdreifachbildungen) in decapod crustacean appendages are described. They are similar to the range of structures experimentally produced by cutting notches in the sides of insect legs (Bohn, 1965). It is argued that they result from failure of wounds to heal. Regeneration from a free surface along the proximodistal axis is always in a distal direction. Surfaces regenerating circumferentially can regenerate in either direction around the circumference. Regeneration will proceed until the two surfaces of the wound meet. Then, where the two surfaces on either side are non-congruent, intervening tissues will be intercalated. This explanation accounts for the range of structures observed after notching experiments (Bohn, 1965) and seen in crustacean Bruchdreifachbildungen. The explanation says that regeneration will occur wherever wounds fail to heal. This avoids the difficulties of the complete circle rule (French, Bryant & Bryant, 1976) and explains why appendages with mirror-image symmetry are often capable of regeneration.