Summary of papers_first week of June

The allometry of algal growth rates. Tang, E.P., 1995.

 allometric relationships for algal growth can be applied to biomass size spectra to yield
estimates of primary productivity.
 Varying scaling exponent of algal growth  reflecting differences in the expression of cell
size or phyletic differences or small sample size.
 127 observations for three algal divisions (Chlorophyta, Chrysophyta, Pyrrophyta) and five
classes (Bacillariophyceae, Chlorophyceae, Chrysophyceae, Dinophyceae and
Prymnesiophyceae)
 Size effect is smaller if algal size is expressed as cell volume, reflecting changes in cell
composition with size.
 Relationship between algal growth rate and cell carbon, u = 3.45C -0.21
o scaling exponent of algal growth is the same for different taxa, but dinoflagellates
tend to have lower growth rates than other algae of similar size.
 When cell size was expressed in terms of volume, the shallower slope of-0.16 was obtained
o in algae, intracellular concentrations of nitrogen, phosphorus and carbon decline with
size  this implies that elemental concentrations change more slowly than volume
o growth rate changes more slowly with cell volume than elemental composition
shallower slope for the algal growth- volume relationship.
 3 divisions did not have significantly different slopes for the growth rate-size curve, but
pyrrophytes, that includes dinoflagellates, have significantly lower growth rates than other
divisions
o In 5 classes, none had significantly different slopes for the growth-size relationship,
but dinoflagellates (i.e. Dinophyceae) have lower growth rates than other classes
o lower photosynthetic capacity per unit biomass of dinoflagellates

Temperature‐and size‐scaling of phytoplankton population growth rates: Reconciling the

Eppley curve and the metabolic theory of ecology
Kremer, C.T., Thomas, M.K. and Litchman, E., 2017

 Strong effects of temperature on population growth rate have inspired two frameworks—
Eppley curve & metabolic theory of ecology—that produce different quantitative
relationships and employ distinct statistical approaches
 Size and functional group membership are also critical.
 Metabolic theory’s temperature-scaling predictions are more accurate
 Eppley curve is a phenomenological description of the temperature- dependence of growth,
whose parameters are estimated from data rather than predicted from first principles.
o based on characterizations of an empirical relationship
o Eppley curve mathematically – u =a exp (b.T), u - maximum growth rate of any
phytoplankton species, which varies with temperature, T. Parameter a (Eppley
component) controls the height of the exponential function at 0 oC while b
determines how strongly u rises with temperature
 MTE offers a contrasting, mechanistic description of the same empirical pattern, derived
from studying the factors constraining individual metabolic rates
o focuses specifically on biochemical reactions essential for life, which depend on
temperature and supply of substrate
o Assuming substrates are not limiting, reaction rates will depend only on
temperature, and can be described by the Arrhenius-van’t Hoff equation: k œ
e-E/RT
o Assumes, rate of the slowest chemical reaction (lowest E) that is essential for life
will determine just how quickly individuals and populations can grow, such that u œ
k
 E & b control the sensitivity of maximum growth rate to temperature - impossible to directly
com- pare these coefficients because the Eppley curve is a function of T, while the
Arrhenius-van’t Hoff equation depends on -1/T
 Increases in population growth rate with temperature are consistent with metabolic theory,
and weaker than previous estimates of the Eppley curve.
o maximum growth rate is limited by the temperature sensitivity of photosynthesis
 A 10 oC temperature increase will increase growth rates by a factor of 1.53, rather than 1.88
as in previous studies of the Eppley curve
 Population growth rates decrease with size, but much less strongly that MTE predicts.
 Growth rates of different functional groups scale similarly with temperature, but some groups
grow faster than others, independent of temperature
o slope of the temperature-scaling relationship is consistent across functional groups
although groups differ in their intercepts (maximum growth rate at T = 0 oC).
 Temperature dependence of growth rate under replete conditions emerges from the shared
biochemistry and thermodynamic constraints of all photoautotrophs.
 diatoms and green algae have intercepts that are higher than the previously estimated
intercept of the Eppley curve
o older parameterizations of the Eppley curve - will underestimate their growth rates
at low temperatures, while overestimating their growth rates at high temperatures
 Heterogeneity in the observed growth rates - a temperature-only model cannot capture, as it
is attributable to functional group identity and cell size.
 Growth rate decreases with cell size, rate of decline was much weaker than MTE predicts (-
0.054 rather than -0.25)
 very little difference in the size-scaling coefficient whether study used ln(biovolume) or
ln(mass), calculating mass using two different conversions
 Larger cells - lower S/V & increased self-shading  growth rate decrease with size, as larger
cells  difficult to get light and nutrient efficiently and distribute internally
o changes in cell shape with increasing size can alleviate these constraints and weaken
the size-scaling relationship
 Small cells may experience limited growth rates because unavoidable investments in non-
scalable nitrogenous compounds such as DNA result in reduced biosynthetic capacity.
 Depending on their strength and shape, combined effects of opposing trade-offs between
being small and large could produce a size-scaling relationship that is weaker than the MTE
proposed -1/4, or even result in a nonlinear relationship
 Both the Eppley and MTE indicated that functional group identity explained significant
variation in the intercept (but not the slopes) of the temperature- and size- scaling
relationships
 Slow growth rate of dinoflagellates than diatoms - dinoflagellates have enormous genomes,
and consistently lower growth rates, ability to both photo- synthesize and consume other
organisms may come at the expense of rapid growth.
 At same temperature, communities dominated by different functional groups will have
substantially different maximum growth rates, and hence, productivities
o While growth rates will increase across temperatures, this hierarchy will not change.

Unimodal size scaling of phytoplankton growth and the size dependence of nutrient uptake
and use. Marañón, E., et al., 2013.

 umax and carbon-specific photosynthesis peak at intermediate cell sizes

 Maximum nitrogen uptake rate (VmaxN) scales isometrically with Vcell, whereas nitrogen
minimum quota scales as Vcell0.84
 Large cells thus possess high ability to take up nitrogen, relative to their requirements, and
large storage capacity, but their growth is limited by the conversion of nutrients into
biomass
 Small species show similar volume specific VmaxN compared to their larger counterparts
but have higher nitrogen requirements.
 unimodal size scaling of phytoplankton growth arises from taxon-independent, size-related
constraints in nutrient uptake, requirement and assimilation.
 unimodal pattern in the relationship between cell size and growth and metabolic rate thus
appears robust, particularly considering the wide cell size and phylogenetic ranges
considered
 As population growth requires the synthesis of new biomass, umax is closely related to
metabolic rate  same unimodal size scaling in carbon-specific CO2 fixation, which
represents a bio- mass turnover rate.
 when a sufficiently large range in body size is considered, a single, scale-free power law
relating metabolic rate with cell size is not applicable to photosynthetic unicells.
 Contrary to photosynthesis, biomass-specific respiration rates did not show any relationship
with cell size.
 respiratory losses represented a small fraction of total carbon fixation, reflecting near-
optimal conditions during the exponential growth phase of cultures
 phytoplankton umax and its size scaling is largely controlled by metabolic gains rather than
losses.
 assuming a constant density of transport sites on the cell membrane, VmaxN should be
linearly related to cell surface area
 VmaxN scales isometrically (slope = 0.97) with cell volume.  rate of nutrient uptake per
unit cell surface area increases with cell size  as a result of an increasing density of
transport sites
 as phytoplankton species become larger, their ability to take up nitrogen, when available in
large concentrations, increases faster than their minimum nitrogen requirement does.
o as cell size becomes smaller, the maximum nutrient uptake rate decreases faster than
the mini- mum nitrogen requirement.
 ability of larger species to sustain high values of VmaxN is associated with an increasing
storage capacity
 higher storage capacity of larger cells - they fill up more slowly when acquiring nutrients 
can sustain high uptake rates for longer periods of time  advantage in environments
where nutrient supply is highly intermittent
 in spite of their high VmaxN and storage capacity, the largest species grew more slowly
than the intermediate size  smaller ability to convert nutrients into biomass
 cell volume increases resources must cover a longer distance from the cell surface to
metabolically processing sites  effect can be partially attenuated by the greater degree of
vacuolation in larger cells
 Reasons for slow growth in larger cells
o Slower nutrient supply
o decrease in the density of enzymatic units slow assimilation of nutrients into biomass slow
o decrease in light absorption growth rate
 small species also showed slower growth rates than intermediate-size cells
 C : N ratio decreased markedly in the species below 50 um3 in cell volume
o increasing relative abundance of nitrogen-containing molecules - nucleic acids and
membrane proteins
o reduced storage of carbon-rich compounds - lipids and carbohydrates.
o smallest species were thus more nitrogen rich, but their VmaxN was, on a volume-
specific basis - similar to that of larger cells
 non-scalable components - membranes and nucleic acids, occupy an increasingly larger
fraction of cell volume as cell size decreases,  a decrease in the fraction of cytoplasm
available for other scalable, catalytic components directly involved in metabolic activity
and biomass production
 decrease of lmax in small cells noticeable below a cell volume of c. 100 um3
 unimodal pattern in umax explain why so many bloom-forming species are of intermediate
cell size