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A
nutrient substrate
gradient of metabolites produced
by the dark cells
substrate
gradient
substrate
B
threshold
nutrient substrate
gradient of metabolites produced
by the dark cells
substrate
gradient
substrate
C
threshold
nutrient substrate
(adapted from Kupiec, 1986, 1997)
F IGURE 16. Model of the heap of cells. The growth of this cell colony depends
on its environment. A: Only the cells of the first layer have direct access to the
substrate. B: The other cells only have access to the substrate which is diffus-
ing in the colony and forming a decreasing concentration gradient, or access to
the metabolites, which form secondary gradients. Below a certain concentration
threshold, the cells can no longer grow (dark cells). C: They must then differ-
entiate to adapt to these microenvironmental conditions and continue to grow
( light cells).
by the concentrations of nutrient and metabolites. The cells situ-
ated at the lower part of the colony have a microenvironment richer
in nutrient while those at the upper part have a microenvironment
richer in metabolites. To optimise their growth, the cells have to
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