PHYSIOLOGICAL RESPONSES OF RHIZOPUS UNDER DIFFERENT ENVIRONMENTAL FACTORS

Rhizopus differs from other molds, because it is non-septate and produces sporangiospores rather than
conidia. It is a very-fast-growing, spreading type of mold which has white mycelia and black sporangia.
Rhizopus species form rhizoids at the base of the sporangiophores, and columella in the sporangium.
Young sporangia are white before turning black with age. Probably the most common of the Rhizopus
species is R. stolonifer, the so-called bread mold.

Besides bread, Rhizopus causes spoilage of strawberries, other berries, fruits, and vegetables. Rhizopus
species have also been isolated from cereal grains, nuts, and meat. R. oligosporus is used in making
tempeh and certain other mold-fermented foods.

Rhizopus is placed in the taxonomic order Mucorales and is somewhat typical, and probably the most
common genus of that order to contaminate foods. Other genera in the order Mucorales which are
nonseptate and produce sporangiospores are Absidia, Mucor, Rhizomucor, Syncephalastrum, and
Thamnidium. All of the Mucorales genera that contaminate food are found in the family Mucoraceae.
These genera are sometimes referred to as mucoraceous fungi. They are a part of the class Zygomycetes
and form zygospores by sexual processes and chlamydospors . They require high-moisture conditions for
growth.

GROWTH OF RHIZOPUS

Rhizopus species grow as filamentous, branching hyphae that generally lack cross-walls (i.e., they are
coenocytic). They reproduce by forming asexual and sexual spores. In asexual reproduction,
sporangiospores are produced inside a spherical structure, the sporangium. Sporangia are supported by
a large apophysate columella atop a long stalk, the sporangiophore. Sporangiophores arise among
distinctive, root-like rhizoids. In sexual reproduction, a dark zygospore is produced at the point where
two compatible mycelia fuse. Upon germination, a zygospore produces colonies that are genetically
different from either parent.

The growth of living organisms is influenced simultaneously by all the components of the environment
and the response to any one environmental factor may be influenced by the levels of other factors. This
observation was first incorporated into a ‘law of tolerance’ by Shelford (1913) and may be stated as
follows: an organism’s tolerance to unfavourable conditions is reduced when other environmental
factors are also at non-optimal levels (Odum 1971; Pianka 1974). Many food preservation processes do,
of course, depend on the use of combinations of environmental factors at unfavourable levels. The
effects on the hyphal extension rate of interactions between environmental factors and the additive
effects of factors at unfavourable values are well illustrated. For example, as water activity is decreased
the hyphal extension rate declines at all values of temperature and pH.

Modification of optimum values by other environmental conditions. The optimum temperature

describes the temperature at which microbes grow most rapidly although it is recognized that, at the
optimum temperature, the organism is to some extent stressed and the accelerating effects of high
temperatures on metabolism are counteracted by degradative reactions. It is commonly observed that
the optimum temperature for microbial growth is lower at unfavourable pH values than at the optimum
pH for an organism (e.g. Chung and Goepfert 1970). This phenomenon is clearly evident with Rhizopus
oligosporus, where the optimum temperature was lowered from ‡ 42°C at pH 5.5 and 7.5 to 36–37°C at
pH 3.5.

The optimum pH for R. oligosporus was also affected by the water activity of the medium, being 5.8 at
water activity of 1.0, 5.6 at 0.98 and 5.5 at 0.96.

The optimum water activity and CO2 concentrations did not appear to be affected by the values of other
environmental factors and remained at 1.0 and 0.03%, respectively, throughout. Rhizopus spp. appear
to be relatively tolerant of reduced aw so long as other conditions are near optimal.

Hocking and Miscamble (1995) examined the effect of a w on the growth of a number of Rhizopus spp.
Rhizopus oryzae and R. microsporus grew equally rapidly at aw 0.96–1.00 while R. stolonifer grew equally
rapidly from 0.90 to1.00. Rhizopus oligosporus appears to be rather more sensitive to low aw than the
Rhizopus species examined by Hocking and Miscamble (1995) and its growth was slower at a w 0.98 than
at1.00. However, the prediction that the hyphal extension rate progressively slows with a decrease in a w
from 1.00 to 0.98 could be a model artefact since no observations were made at intermediate a w.