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differential adhesion hypothesis
Differential adhesion hypothesis (DAH) is a hypothesis that explains cellular movement during morphogenesis with thermodynamic principles. In DAH tissues are treated as liquids consisting of mobile cells whose varying degrees of surface adhesion cause them to reorganize spontaneously to minimize their interfacial free energy. Put another way, according to DAH, cells move to be near other cells of similar adhesive strength in order to maximize the bonding strength between cells and produce a more thermodynamically stable structure. In this way the movement of cells during tissue formation, according to DAH, parodies the behavior of a mixture of liquids. Although originally motivated by the problem of understanding cell sorting behavior in vertebrate embryos, DAH has subsequently been applied to explain several other morphogenic phenomena.
==Background==
The origins of DAH can be traced back to a 1955 study by Philip Townes and Johannes Holtfreter.〔 In this study Townes and Holtfreter placed the three germ layers of an amphibian into an alkaline solution, allowing them to dissociate into individual cells, and mixed these different types of cells together. Cells of different species were used to be able to visually observe and follow their movements. Cells of similar types migrated to their correct location and reaggregated to form germ layers in their developmentally correct positions. This experiment demonstrated that tissue organization can occur independent of the path taken, implying that it is mediated by forces that are persistently present and doesn't arise solely from the chronological sequence of developmental events preceding it.〔
From these results Holtfreter developed his concept of selective affinity, and hypothesized that well-timed changes to selective affinity of cells to one another throughout development guided morphogenesis.〔 Several hypotheses were introduced to explain these results including the "timing hypothesis" and the "differential surface contraction hypothesis".〔 In 1964 Malcolm Steinberg introduced the "differential adhesion hypothesis" which uses thermodynamic principles to describe and explain patterns of cell sorting and arrangement observed.〔
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