Biological Physics of the Developing Embryo

	Acknowledgments	page vii
	Introduction: Biology and physics	1
1	The cell: fundamental unit of developmental systems	6
	The eukaryotic cell	6
	Diffusion	8
	Osmosis	15
	Viscosity	16
	Elasticity and viscoelasticity	21
	Perspective	22
2	Cleavage and blastula formation	24
	The cell biology of early cleavage and blastula formation	24
	Physical processes in the cleaving blastula	29
	Physical models of cleavage and blastula formation	39
	Perspective	50
3	Cell states: stability, oscillation, differentiation	51
	Gene expression and biochemical state	52
	How physics describes the behavior of a complex system	53
	Oscillatory processes in early development	57
	Multistability in cell-type diversification	63
	Perspective	76
4	Cell adhesion, compartmentalization, and lumen formation	77
	Adhesion and differential adhesion in development	78
	The cell surface	80
	Cell adhesion: specific and nonspecific aspects	81
	The kinetics of cell adhesion	84
	Differential adhesion of embryonic tissues	90
	The physics of cell sorting	95
	Perspective	97
5	Epithelial morphogenesis: gastrulation and neurulation	99
	Physical properties of epithelia	100
	Gastrulation	108
	Convergence and extension	117
	Neurulation	122
	Perspective	128
	Appendix: Linear stability analysis	128
6	Mesenchymal morphogenesis	131
	Development of the neural crest	134
	The extracellular matrix: networks and phase transformations	138
	Mesenchymal condensation	149
	Perspective	154
7	Pattern formation: segmentation, axes, and asymmetry	155
	Basic mechanisms of cell pattern formation	157
	Segmentation	162
	Epithelial patterning by juxtacrine signaling	168
	Mesoderm induction by diffusion gradients	171
	Reaction–diffusion systems	173
	Control of axis formation and left–right asymmetry	177
	Perspective	187
8	Organogenesis	188
	Development of the cardiovascular system	190
	Fractals and their biological significance	197
	Branching morphogenesis: development of the salivary gland	203
	Vertebrate limb development	210
	Perspective	222
9	Fertilization: generating one living dynamical system from two	223
	Development of the egg and sperm	224
	Interaction of the egg and sperm	233
	Propagation of calcium waves: spatiotemporal encoding of postfertilization events	236
	Surface contraction waves and the initiation of development	242
	Perspective	247
10	Evolution of developmental mechanisms	248
	The physical origins of developmental systems	249
	Analyzing an evolutionary transition using physical concepts: segmentation in insects	256
	The evolution of developmental robustness	262
	Perspective	272
	Glossary	273
	References	291
	Index	327

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Gabor Forgacs

University of Missouri, Columbia

Stuart A. Newman

New York Medical College

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