In
his research Max Tilzer has been engaged mainly in work on production
biology with emphasis on the phytoplankton, both in freshwater lakes
and in the ocean. His lake research has been conducted in Vorderer Finstertaler
See, Austria, a small high-mountain lake, large and extremely clear
Lake Tahoe, USA, and in mesotrophic Lake Constance, Germany-Switzerland-Austria.
A major research topic has been the utilization of underwater light
for the primary production process, and the role of primary production
in controlling the dynamics of the phytoplankton community. His oceanographic
research mainly was conducted in the Southern Ocean where he was focusing
on the effects of the prevailing low water temperatures on the primary
production process. Moreover, he participated in research in the Gulf
of Aqaba / Red Sea.
Max
Tilzer could show that, as a general rule, aquatic ecosystems are less
productive than terrestrial ones. Main reasons for this is that water
as a medium is considerably more opaque than air. As a consequence,
aquatic phototrophic organisms have to compete for photons with the
medium surrounding them. By contrast, closed terrestrial plant stands
are able to absorb and subsequently utilize virtually all light impinging
on the Earth surface. In dense phytoplankton, water transparency is
diminished by phytoplankton self-shading, thus reducing the vertical
extent of the productive layer. Due to self-shading, areal production
increases with biomass according to a saturation hyperbola. This saturation
curve can also be explained by the fractional light absorption due to
phytoplankton with increasing pigment concentration relative to background
absorption by water. The theoretical upper limit of aquatic primary
productivity would be reached if 100% of the light penetrating the water
would be absorbed by the photosynthetic pigments. As a rule, however,
only a relatively small proportion of the light entering the water is
actually absorbed by plant pigments.
Because, moreover, even clear water strongly absorbs red light, only
the blue-light absorption maximum of aquatic phototrophs can be utilized
for photosynthesis. By deep vertical water column mixing, the light
available to phytoplankton is further diminished because algae travel
through a light gradient, together with the circulating water masses.
Surface inhibition, especially in clear waters, reduces photosynthesis
further. By acclimation to the prevailing light climate, phytoplankton
can to some extent increase the utilization efficiency of underwater
light for pghotosynthesis. The most common mechanism consists of alterations
in the cellular pigment contents. When the productive layer is thermally
stratified, surface algae are acclimated to higher radiances than phytoplankton
in deeper water. Another strategy to optimize the light environment,
consists of diel vertical migrations of flagellated phytoplankton.
The quantum yield of photosynthesis reaches its highest value only in
deeper water where light is limiting. The highest values reached are
0.03 - 0.04, which is considerably below the theoretical upper limit
of the quantum yield of 0.125. For all the above-mentioned reasons,
the overall light utilization efficiency by phytoplankton photosynthesis
is generally low. Only in exceptional cases with high phytoplankton
biomass, more than 1% of the incident short-wave radiant energy is converted
into chemically stored energy in photosynthates.
Investigations
in Vorderer Finstertaler See and Lake Constance revealed that the relationship
between areal daily gross primary production rates and rates of biomass
increase is poor. This can be explained by the high proportions of metabolic
(mainly respiratory) losses relative to gross photosynthesis during
the primary production process. Moreover, metabolic losses are highly
correlated with potential biomass growth rates as derived from photosynthesis
estimate. The high proportions of metabolic losses during the primary
production process restricts the transfer efficiency of energy and organic
carbon from the primary producers to the subsequent food web.
The
dramatic decrease of external phosphorus loading of Lake Constance during
the 1980’s, as a consequence of sewage treatment and wastewater
diversion, did not immediately result in decreasing annual primary productivity.
It was hypothesized that during the ”eutrophic” phase of
Lake Constance, intense grazing by zooplankton as well as sedimentation
led to an effective removal of phytoplankton from the euphotic zone,
thus preventing the nutrient-dependant carrying capacity to be reached.
As phosphorus loading decreased, the annual biomass initially remained
unchanged thus increasing the nutrient-dependent carrying capacity.
Photosynthetic
measurements during the first Antarctic expedition of Max Tilzer showed
that chlorophyll-specific photosynthetic rates, both at saturating and
at limiting light levels, where significantly smaller than in warmer
environments. Subsequent laboratory experiments on board of ”Polarstern”
on the temperature-dependence of photosynthesis suggested that chlorophyll-specific
photosynthetic rates were temperature-sensitive both at saturating and
at limiting light levels. In another study, the temperature dependence
of light-saturated photosynthesis and respiration were compared. It
could be shown that respiration tends to be more strongly diminished
at low temperatures than is photosynthesis. This would mean that at
extremely low water temperatures, the mass balance between photosynthesis
and respiration is shifted in favor of photosynthesis. Model calculations
based on these experimental results have suggested that algal population
growth rates are not as much decreased by low temperatures than one
would expect from experience at high temperatures. This could be of
particular importance during seasons with short daylengths when nightly
respiratory carbon losses are comparatively small.
Extensive
studies on the spectral composition of the underwater light of the Southern
Ocean have revealed the extreme water clarity of this predominantly
unproductive sea. The main reason for the exceptionally low non-algal
vertical light attenuation in Antarctic waters is the minimal terrestrial
influence. Because in most cases blue light is best transmitted, light
absorption cross-sections of the phytoplankton cell suspensions are
comparatively high as compared to green and/or turbid waters. As a consequence,
phytoplankton has a comparatively strong influence on both overall light
attenuation and the spectral composition of underwater light.
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