Biofilms: Microbial Life on Surfaces

Rodney M. Donlan


Emerging Infectious Diseases. 2002;8(9) 

In This Article


Biofilm cells may be dispersed either by shedding of daughter cells from actively growing cells, detachment as a result of nutrient levels or quorum sensing, or shearing of biofilm aggregates (continuous removal of small portions of the biofilm) because of flow effects.

The mechanisms underlying the process of shedding by actively growing cells in a biofilm are not well understood. Gilbert et al.[69] showed that surface hydrophobicity characteristics of newly divided daughter cells spontaneously dispersed from either E. coli or P. aeruginosa biofilms differ substantially from those of either chemostat-intact biofilms or resuspended biofilm cells. These researchers suggested that these differences might explain newly divided daughter cells' detachment. Hydrophobicity was lowest for the newly dispersed cells and steadily increases upon continued incubation and growth.

Alginate is the major component of the EPS of P. aeruginosa. Boyd and Chakrabarty[70] studied alginate lyase production in P. aeruginosa to determine whether increased expression of this enzyme affected the size of the alginate molecules (and therefore adhesion of the organisms). Inducing alginate lyase expression substantially decreased the amount of alginate produced, which corresponded with a significant increase in the number of detached cells. The authors suggested that the role of algL (the gene cassette for alginate lyase production) in wild type P. aeruginosa may be to cause a release of cells from solid surfaces or biofilms, aiding in the dispersal of these organisms. Polysaccharidase enzymes specific for the EPS of different organisms may possibly be produced during different phases of biofilm growth of these organisms.

Detachment caused by physical forces has been studied in greater detail. Brading et al.[71] have emphasized the importance of physical forces in detachment, stating that the three main processes for detachment are erosion or shearing (continuous removal of small portions of the biofilm), sloughing (rapid and massive removal), and abrasion (detachment due to collision of particles from the bulk fluid with the biofilm). Characklis[72] noted that the rate of erosion from the biofilm increases with increase in biofilm thickness and fluid shear at the biofilm-bulk liquid interface. With increase in flow velocity, the hydrodynamic boundary layer decreases, resulting in mixing and turbulence closer to the biofilm surface. Sloughing is more random than erosion and is thought to result from nutrient or oxygen depletion within the biofilm structure.[71] Sloughing is more commonly observed with thicker biofilms that have developed in nutrient-rich environments.[72] Biofilms in fluidized beds, filters, and particle-laden environments (surface waters) may be subject to abrasion.

Detachment is probably also species specific; P. fluorescens disperses and recolonizes a surface (in a flow cell) after approximately 5 h, V. parahaemolyticus after 4 h, and V. harveyi after only 2 h.[73] This process probably provides a mechanism for cells to migrate from heavily colonized areas that have been depleted of surface-adsorbed nutrients to areas more supportive of growth.

The mode of dispersal apparently affects the phenotypic characteristics of the organisms. Eroded or sloughed aggregates from the biofilm are likely to retain certain biofilm characteristics, such as antimicrobial resistance properties, whereas cells that have been shed as a result of growth may revert quickly to the planktonic phenotype.