Microbial Extracellular Enzyme Activity: A New Key Parameter Aquatic Ecology

4
Microbial Extracellular
Enzyme Activity: A New Key
Parameter in Aquatic Ecology
Hans-Georg Hoppe
4.1 Introduction
Three general pathways of organic matter degradation exist in natural aquatic

environments. These are based on predation, particle feeding, and dissolved
organic matter (DOM) uptake. Bacteria are involved in the latter two in that
they are able to hydrolyze nonliving particles thereby competing with particle
feeders, and take up small organic molecules, which is their exclusive domain.
Particle hydrolysis is mediated by extracellular enzymes in the intestines of an-
imals and by the enzymatic activity of attached bacteria. Therefore, successful
competition for organic matter among tropic levels is also a question of extra-
cellular enzymatic efficiencies. The decomposition of dissolved organic mac-
romolecules is mediated mainly by the enzymes of free-living bacteria, which
subsequently incorporate the small molecules resulting from enzymatic hydro-
lysis. Therefore, bacterial activity has a strong influence on the concentration
and speciation of dissolved organic molecules in the water. A major fraction of
the DOM-pool in the water can be expected to consist of dissolved macro-
molecules since extracellular hydrolysis is a relatively slow process in compar-
ison to the uptake of low-molecular-weight organic matter (LMWOM). The ef-
ficiency of animals feeding on particles may vary considerably, depending on
various factors. Microbial hydrolysis of particles will depend greatly on the
chemical composition and the size of the particles. Thus competition for organic
particles between animals and microbes will be determined on the one hand,
by the slow but continuous microbial component and, on the other hand, by
pulse-feeding activities of animals ijoint and Morris, 1982). In detail, it has been
pointed out that efficiencies of microbial decomposition of fast-sinking large
60
R. J. Chróst (ed.), Microbial Enzymes in Aquatic Environments
© Springer-Verlag New York Inc. 1991
Chapter 4 Microbial Extracellular Enzyme Activity 61
particles and nonsinking small particles may be different (Cho and Azam, 1988;
Karl et aI., 1988).
The input of substances into the pool of DOM is, by necessity, mediated
by the hydrolytic activity of bacteria attached to particles and, to a certain de-
gree, by autolysis of cells, sloppy feeding, and excretion of plants and animals.
The attached bacteria, however, contribute to the DOM pool and benefit the
free-living bacteria only when they do not entirely use up the products of par-
ticle hydrolysis themselves. Overproduction of hydrolysis products, as dem-
onstrated in this report, causes a considerable shift in organic materials from
the particulate to the dissolved state. This process is mediated by the different
enzymatic strategies of attached and free-living microorganisms and provides
the basis for their nutritional demands. Therefore, it is suggested that the mi-
crobialloop model (Azam et aI., 1983) be amended to include the initial and
obligatory hydrolytic steps of microbial nutrient generation, which would serve
to set the range of model dimensions (Chr6st, 1990). The microbial loop model
(Figure 4.1) describes the role of bacteria and bacteriovorous organisms in or-
ganic matter transfer and mineralization, these processes, however, strongly
depend on the enzymatic capacities of the microbial community.
Microbial attachment is a broad field of research, many aspects of which
have already been investigated, ranging from adhesion mechanisms to meta-
bolic properties of attached bacteria (Hoppe, 1984) and their chemotactic re-
sponse Gackson, 1989). However, information on enzymatic activity of bacteria
on surface materials is scarce, despite the fact that it is probably one of the
most important functions of surfaces in the aquatic environment. Therefore it
I microbial loop II
primary
- exudates - autolysis
producers
living deL dissolved
J +
animals~detntus~EEA~
POlyi,ers
I'
olgomers
~EEA~monomers
I
uP~ke
bacteria
het. flagellates -ciliates
biomass+
respiration
Figure 4.1 Bacterial loop model supplemented with pathways of organic matter trans-
formation from primary producers to bacteria via extracellular enzymatic activity (EEA).

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Three general pathways of organic matter degradation exist in natural aquatic

living deL dissolved

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