In evolutionary biology, robustness of a biological system (also called biological or genetic robustness) is the persistence of a certain characteristic or trait in a system under perturbations or conditions of uncertainty. Robustness in development is known as canalization. According to the kind of perturbation involved, robustness can be classified as mutational, environmental, recombinational, or behavioral robustness etc. Robustness is achieved through the combination of many genetic and molecular mechanisms and can evolve by either direct or indirect selection. Several model systems have been developed to experimentally study robustness and its evolutionary consequences.
Classification
Mutational robustness
Mutational robustness (also called mutation tolerance) describes the extent to which an organism's phenotype remains constant in spite of mutation. Robustness can be empirically measured for several genomes and individual genes by inducing mutations and measuring what proportion of mutants retain the same phenotype, function or fitness. More generally robustness corresponds to the neutral band in the distribution of fitness effects of mutation (i.e. the frequencies of different fitnesses of mutants). Proteins so far investigated have shown a tolerance to mutations of roughly 66% (i.e. two thirds of mutations are neutral).
Conversely, measured mutational robustnesses of organisms vary widely. For example, >95% of point mutations in C. elegans have no detectable effect and even 90% of single gene knockouts in E. coli are non-lethal. Viruses, however, only tolerate 20-40% of mutations and hence are much more sensitive to mutation.
Robustness to stochasticity
Biological processes at the molecular scale are inherently stochastic. They emerge from a combination of stochastic events that happen given the physico-chemical properties of molecules. For instance, gene expression is intrinsically noisy. This means that two cells in exactly identical regulatory states will exhibit different mRNA contents. The cell population level log-normal distribution of mRNA content follows directly from the application of the Central Limit Theorem to the multi-step nature of gene expression regulation.
Environmental robustness
In varying environments, perfect adaptation to one condition may come at the expense of adaptation to another. Consequently, the total selection pressure on an organism is the average selection across all environments weighted by the percentage time spent in that environment. Variable environment can therefore select for environmental robustness where organisms can function across a wide range of conditions with little change in phenotype or fitness (biology). Some organisms show adaptations to tolerate large changes in temperature, water availability, salinity or food availability. Plants, in particular, are unable to move when the environment changes and so show a range of mechanisms for achieving environmental robustness. Similarly, this can be seen in proteins as tolerance to a wide range of solvents, ion concentrations or temperatures.
Genetic, molecular and cellular causes
Genomes mutate by environmental damage and imperfect replication, yet they display remarkable tolerance. This comes from robustness both at many different levels.
Organism mutational robustness
There are many mechanisms that provide genome robustness. For example, genetic redundancy reduces the effect of mutations in any one copy of a multi-copy gene. Additionally the flux through a metabolic pathway is typically limited by only a few of the steps, meaning that changes in function of many of the enzymes have little effect on fitness. Similarly metabolic networks have multiple alternate pathways to produce many key metabolites.