Diatom–environmental relationships and nutrient transfer functions from contrasting shallow and deep limestone lakes in Ontario, Canada

Diatom assemblages were analysed in the surface sediments of 44 alkaline lakes in south-western Ontario, Canada, and combined with a pre-existing 58 south-eastern Ontario lake set: (1) to determine if shallow, polymictic Ontario lakes contain different diatom assemblages from deeper, dimictic lakes, and if so, which environmental variables most influence assemblages; (2) to improve the existing transfer functions; (3) to construct and compare transfer functions separately for dimictic, deep lakes and for polymictic, shallow lakes. Polymictic and dimictic lakes covered a similar nutrient range (spring total phosphorus (TP)=4–54 g/l, spring total nitrogen (TN)=200–927 g/l; n=101) and spring pH levels (7.6–9; n=101). However, polymictic lakes were shallower (median mean depth = 2.9 m vs. 7.3 m in dimictic lakes). Benthic diatoms (average 60% relative abundance) dominated the polymictic lakes, whereas planktonic diatoms (average 60%) dominated dimictic lakes. A Canonical Correspondence Analyses with forward selection (p < 0.05, 999 Monte Carlo permutations) identified TP, alkalinity, watershed to volume ratios and lake depth as the most important measured environmental variables influencing diatom distribution in both polymictic and dimictic lakes. Additionally, pH was identified as an important variable in polymictic lakes, whereas TN was also forward selected in the dimictic lakes. Adding more lakes to the original southern Ontario calibration set improved the TN transfer function (r² _jack=0.42, root mean squared error of prediction (RMSEP)_jack=0.11 [log g TN/l]), although there was a high systematic error in the revised model (r² _residual = 0.48). However, the strongest TP model was derived from the polymictic lakes (r² _boot =0.44, RMSEP_boot=0.20 [log g TP/l]), which was the smallest lake set (n=30) with the lowest number of diatom species. The stronger TP model from the polymictic lakes may be partly due to the relatively low macrophyte cover in our polymictic lakes, which may lead to stronger benthic–pelagic coupling than in lakes with large macrophyte populations. Additionally, our study suggests that the Chrysophyceae cyst:diatom frustule ratio may be useful for indicating trends in TP levels of 35 g/l in alkaline lakes that are dimictic, but is not necessarily indicative of trophic state changes in shallow, polymictic lakes. Our study demonstrates that it may be important to construct separate diatom-based nutrient transfer functions for polymictic and dimictic lakes.     

Adaptation of inducible defense in Euplotes daidaleos (Ciliophora) to predation risks by various predators

The extent of induced morphological defense in Euplotes daidaleos correlates to this ciliate's predation risk from the defense-inducing predator species. Euplotes daidaleos responded by morphological transformation only to organisms that are able to feed on typically formed Euplotes cells (63 ± 5 m cell width in E. daidaleos). Three of those potential predator species caused defensive changes to various degrees (Student's t-test, P < 0.1 to P < 0.0001): Lembadion bullinum (Ciliata) induced 82 ± 6 m cell width in E. daidaleos; Chaetogaster diastrophus (Oligochaeta) induced 85 = 6 m width; and Stenostomum sphagnetorum (Turbellaria) induced 89 ± 8 m width (at a density of 10 predators per milliliter, respectively). At higher predator densities (50 or 100 organisms per milliliter), Euplotes developed a correspondingly larger width (to a maximum of 103 ± 10 m in the presence of S. sphagnetorum). Euplotes did not respond to organisms (e.g., Blepharisma japonicum, Colpidium campylum, Didinium nasutum, Paramecium caudatum, Spirostomum ambiguum, Stentor coeruleus) that cannot feed on this ciliate species. Daphnia longispina and Bursaria truncatella predators, which can feed on large prey of 125, or 200 m in diameter, respectively, also had no effect on the morphology of Euplotes. The extent of defense in Euplotes that was induced by 10 predators per milliliter during 24 h decreased the predation risk from those predators to 67% in the presence of S. sphagnetorum, to 50% with L. bullinum, and to 15% with C. diastrophus, compared to the typical form of Euplotes. In a natural population, the defensive form of E. daidaleos was found with average cell widths of 88 ± 8 m. The results indicate that predator-induced defense in natural Euplotes populations is beneficial to this prey and that it is adapted to the predation abilities of Euplotes predators, whereby energetical costs related to defensive changes may be saved.     

Vertical distribution of planktonic ciliates in strongly stratified temperate lakes

The vertical distribution of planktonic ciliates in eight strongly stratified temperate lakes was studied in summer 1998. Ciliate abundance and biomass were highest (mean 39.9 cells ml^–1 and 181.9 g C l^–1) in the epi-, and lowest (mean 8.2 cells ml^–1 and 97.6 g C l^–1) in the hypolimnion. The community of ciliates was dominated by five orders: Oligotrichida, Haptorida, Prostomatida, Scuticociliatida and Peritrichida. The community composition varied greatly with depth. In the epilimnion, the ciliate numbers were dominated by oligotrichs but small algivorous prostomatids, peritrichs and haptorids were also numerous. In the metalimnion, these groups were replaced by scuticociliates and mixotrophic prostomatids. In the hypolimnion species known as benthic migrants appeared. We found a positive significant correlation (p < 0.05) between ciliate numbers and Chl a and bacterial densities. Only in the hypolimnion, the correlation between ciliates numbers and Chl a was not significant.     

Arsenic concentrations in water and fish from Chautauqua Lake,New York

Synopsis Arsenic persists in Chautauqua Lake, New York waters 13 years after cessation of herbicide (sodium arsenite) application and continues to cycle within the lake. Arsenic concentrations in lake water ranged from 22.4–114.81 g l^–1, = 49.0 ag l^–1. Well water samples generally contained less than 10 g l^–1 arsenic. Arsenic concentrations in lake water exceeded U.S. Public Health Service recommended maximum concentrations (10 g l^–1) and many samples exceeded the maximum permissible limit (50 g l^–1). Fish accumulated arsenic from water but did not magnify it. Fish to water arsenic ratios ranged from 0.4–41.6. Black crappie (Pomoxis nigromaculatus) contained the highest arsenic concentrations (0.14–2.04 g g^–1 ), X = 0.7 g g^–1) while perch (Perca flavescens), muskellunge (Esox masquinongy) and largemouth bass (Micropterus salmoides) contained the lowest concentrations (0.02–0.13 g g^–1). Arsenic concentrations in fish do not appear to pose a health hazard for human consumers.     

A comparative study on planktonic ciliates in two shallow mesotrophic lakes (China): species composition, distribution and quantitative importance

The species richness and seasonal development of planktonic ciliates were studied and compared in two shallow mesotrophic lakes, one covered with dense submerged macrophytes, the other macrophyte poor. Considerable differences in ciliate species composition, dominant taxa, abundance and biomass were observed. Ciliates were much more species rich in the macrophyte-rich lake, while they were more abundant numerically in the macrophyte-poor lake. Altogether, 96 species, included in 53 genera, 14 orders were identified. Among them, 80 species (included in 45 genera, 14 orders) observed from the macrophyte-rich lake, against 49 species (36 genera, 12 orders) were from the macrophyte-poor lake. In the macrophyte-rich lake, the mean abundance and biomass were 13.5 cells ml^-1 and 547.10 g l^-1 f.w.; abundance and biomass were higher in spring and winter; naked oligotrichs dominated total ciliate abundance and Peritrichida dominated the biomass. In the macrophyte-poor lake, ciliate mean abundance and biomass were 35.5 cells ml^-1 and 953.39 g l^-1 f.w.; abundances peaked in autumn; Scuticociliates dominated the abundance and Tintinnids dominated the biomass. Possible causes for the observed differences are discussed.     

What factors control planktonic ciliates during summer in a highly eutrophic lake?

A mesocosm experiment in 24 enclosures (6 m³) started at the end of June 1996 in a highly eutrophic shallow lake, Lake Köyliönjärvi (SW Finland). The original factorial design with nutrient, fish and macrophyte treatments was lost due to strong winds causing leakages. However, after the walls were made leak-proof again on July 11, the planktonic communities developed in divergent ways. On July 31 there was a tenfold variation in total crustacean biomass among the enclosures and the lake (40.2–417.5 g C l^–1), and chlorophyll a varied from 9.5 to 67.0 g l^–1. Here, the single-day data on the 25 planktonic communities is analysed by means of correlation and factor analysis in order to identify factors controlling the protozoans, with particular emphasis on ciliates. The data set comprised: total phosphorus, nitrogen, chlorophyll, bacteria, autotrophic picoplankton, heterotrophic flagellates, abundance and species composition of ciliates, phytoplankton and metazooplankton. The results indicate that although the total ciliate abundance (ranging from 16.2 to 95.0 ind l^–1) was controlled by food resources, the observed differences in ciliate community structure could be attributed partly to differential predation by metazooplankton. The effect of Daphnia cucullata, the dominant daphnid cladoceran, was stronger than that of other metazoans.     

Analysis of the community structure of planktonic ciliated protozoa relative to trophic state in Florida lakes

The planktonic ciliate populations of 30 Florida lakes constituting a broad trophic gradient were examined to determine the response of protozoan community structure to increasing eutrophication. Both ciliate abundance and biomass were strongly related to lake trophic state. Comparison of the Florida data base with a comparable north temperate lake group indicated that subtropical lakes generally possess higher ciliate abundance and biomass at a given trophic state than temperate lakes. However the equations derived for each data base were not significantly different. Community diversity and species richness increased with increasing lake productivity. Highly acidic lakes displayed significantly reduced diversity and numbers of species when contrasted with nonacidic oligotrophic lakes. Small-bodied (< 30 um) ciliates dominated all lakes but were proportionally less important in oligotrophic lakes. Presence-absence data produced three assemblages: an ubiquitous association of primarily small ciliate taxa, a group of large ciliates mainly restricted to eutrophic-hypereutrophic lakes, and a very large ciliate,Stentor niger, which dominated the protozoan communities of acidic oligotrophic lakes.     

The distribution of microplankton in the McMurdo Dry Valley Lakes,Antarctica: response to ecosystem legacy or present-day climatic controls?

Plankton abundance and biomass were investigated in five lakes of the McMurdo Dry Valleys, Antarctica: Lakes Bonney, Fryxell, Joyce, Hoare and Miers. Despite plankton communities being dominated by organisms <100 m in length, there were striking differences between the lakes, including large variations in plankton vertical distribution and differences in total plankton biomass. Bacterial biomass was highest in the anoxic monimolimnia of the meromictic lakes, reaching 191 g C l^–1 in Lake Fryxell. Photosynthetic nanoflagellates dominated phytoplankton in the five lakes studied. Highest chlorophyll a concentrations were recorded at the chemocline of Lake Fryxell (21 g chl a l^–1). Heterotrophic nanoflagellate concentrations were low, ranging from 2 cells ml^–1 in Hoare to 237 cells ml^–1 in Bonney. By Antarctic standards, ciliates were relatively successful in terms of biomass and diversity in Lakes Fryxell and Hoare. In contrast, Lake Miers possessed extremely low ciliate abundance (<0.04 cells ml^–1). On both sampling occasions, copepod nauplii were observed in Lake Joyce. This is the first recording of crustacean zooplankton within the McMurdo Dry Valley Lakes. Because the foodwebs of these lakes are structured by bottom-up forces, differences in plankton distributions could be related to the physicochemical characteristics of each lake. The effect of lake evolution (legacy) and present-day climate change on planktonic dynamics is discussed.     

Nutrient enrichment and nutrient regeneration stimulate bacterioplankton growth

Bacterial abundance results from predatory losses of individuals and replacement of losses through growth. Growth depends on sustained input of organic substrates and mineral nutrients. In this work we tested the hypothesis that bacterial growth in two oligotrophic Canadian shield lakes was limited by nitrogen (N) or phosphorus (P). We also determined whether consumer-regenerated resources contributed substantially to net bacterial growth. Two types of dilution assays were conducted to determine the response of bacteria to nutrient enrichment: diluted whole water (DWW, 1:9 whole/filtered with 0.2 m of filtered lake water) and diluted fractionated water (DFW, 1.0 m prefiltered then diluted as above). Replicate bottles in each dilution assay received either N (50 m), P (10 m), or both N and P enrichments. Controls received no nutrients. Resource-saturated growth rates and grazing rates were estimated from a standard dilution-growth approach. Bacterial growth was stimulated by addition of P alone and in combination with N. Consumers regenerated sufficient resources to support up to half the bacterial growth rate, but the benefit derived from consumers was minor when compared to mortality.     

The phytoplankton of Lake Liddell,New South Wales: chlorophyll a concentrations,species seasonal succession,and covariation with nutrients

Samples of the phytoplankton in a freshwater lake, Lake Liddell, New South Wales (Lat: 32° 22 S, Long. 150° 1 E) were collected every 4 weeks between October 1987 and November 1988. Chlorophyll a concentrations ranged from 1.8 g 1^–1 to 9.1 g 1^–1 and were positively correlated with the following nutrient parameters: total and nett mass additions of nitrate/nitrite-N and total-N, total additions of Kjeldahl-N, and nett mass addition N-P ratios. There was no correlation between lake nutrient concentrations and chlorophyll a. Factors other than nutrient concentrations appeared to be effecting chlorophyll a concentrations as summer levels were low despite nutrient concentrations being at a maximum. In spring and summer the phytoplankton was dominated by chlorophytes, with dinoflagellates and diatoms most abundant in autumn. During winter cyanobacteria were the most abundant. The relative abundance of chlorophytes was positively correlated with in lake nitrate/nitrite-N concentrations whereas the relative abundance of cyanobacteria was negatively correlated with this parameter. Based on chlorophyll a concentrations and the phytoplankton flora Lake Liddell can be classified as mesotrophic.     

Influence of phosphate and nitrate supply on root hair formation of rape,spinach and tomato plants

Summary Experiments with tomato, rape and spinach in nutrient solutions have shown that the formation of root hairs is strongly influenced by phosphate and nitrate supply. Decreasing the phosphate concentration of the nutrient solution from 100 to 2 M P resulted in an increase of root hair length from 0.1–0.2 to 0.7 mm of the three plant species. Root hair density also increased by a factor of 2–4 when the P concentration was lowered from 1000 to 2 M. The variation of these two root properties raised the root surface area by a factor of 2 or 3 compared to plants well supplied with P. Root hair length was closely related to the phosphate content of the root and shoot material. On the other hand, spinach plants grown in a split-root experiment produced root hairs in solutions of high P concentration (1000M P) if the major part of the total root system was exposed to low P concentration (2 M P). It is therefore concluded that the formation of root hairs does not depend on directly the P concentration at the root surface but on the P content of the plant.Similar experiments with nitrate also resulted in an increase in length and density of root hairs with the decrease of concentration below 1000 M. In this case marked differences between plant species occurred. At 2 M compared to 1000 M NO₃ root hair length of tomato increased by a factor of 2, of rape by a factor of 5 and of spinach by a factor of 9. Root hair length was correlated, but not very closely, to the total nitrogen content of the plants. It is concluded, that the influence of nutrient supply on the formation of root hairs is a mechanism for regulating the nutrient uptake of plants.Dedicated to Prof. Dr. E. Welte on the occasion of his 70th anniversary.     

Grazing by a dominant rotifer Conochilus unicornis Rousselet in a mountain lake: in situ measurements with synthetic microspheres

Grazing rates of zooplankton were analysed in the summer of 1999 in Yellow Belly Lake, an oligotrophic system in the Sawtooth Mountains of Idaho (U.S.A.). The colonial rotifer Conochilus unicornis was a dominant species in the epilimnion, with densities reaching 20 colonies l^–1 (ca. 400 ind. l^–1). Clearance rates were measured with an in situ Haney Grazing chamber and synthetic microspheres 5, 9 and 23m in diameter. At epilimnetic temperatures of around 14 °C, mean clearance rates for 5m particles ranged from 30 to 65 l ind.^–1 h ^–1. Clearance rates were 2–9 times higher on the 5m spheres than on the 9 m spheres, and C. unicornis almost never fed on the 23 m spheres. Grazing rates did not change over the diel cycle. Clearance rates declined more than 10-fold as temperatures declined from 14 °C in the epilimnion to 7 °C in the metalimnion. In the epilimnion, grazing by C. unicornis was more important than grazing by crustaceans in the community, at least on particles 9m. The results show the importance of grazing by rotifers in lakes, and the significance of spatial variations that influence grazing rates.     

Clearance rates of bacteria-sized particles by freshwater ciliates,measured with monodisperse fluorescent latex beads

Summary Monodisperse fluorescent latex particles with diameters of 0.57 and 1.04 m have been used to measure the clearance rates of bacteria-sized particles by two ciliates from a Norwegian lake. The clearance rates by Epistylis rotans on these particles were in the range 0.23 to 1.26 l ind^-1h^-1, and by Strombidium sp. 0.26 to 0.90 l ind^-1 h^-1. The clearance rates varied with food level and temperature. Possibilities and limitations of the suggested method are discussed.     

In vitro propagation of Asparagus maritimus – A rare Mediterranean salt-resistant species

Asparagus maritimus L. Miller is a rare species growing of the Mediterranean region and is morphologically similar to A. officinalis. In order to establish an efficient in vitro propagation protocol, explants were excised from spear segments and cultured on Murashige and Skoog (1962) medium containing 3% sucrose and various concentrations of growth regulators. The best shoot initiation (3–4 per explant) was achieved on a medium containing 0.88 M N⁶-benzyladenine (BA), 0.93 M kinetin, 1.07 M -naphthaleneacetic acid (NAA) and 3.90 M ancymidol. Shoot initiation could also be achieved without ancymidol but the shoots were thinner and longer. A very high shoot multiplication rate was achieved on media supplemented with 3% sucrose, 1.07 M NAA, 0.93 M kinetin, 0.44 M BA and various concentrations of ancymidol. The lowest concentration of ancymidol (0.39 M) significantly promoted the highest shoot multiplication rate (11.9 shoots/crown). For root formation, media were supplemented with 6% sucrose, 1.07 M NAA and various concentrations of ancymidol. Rooting frequency increased with higher ancymidol concentration up to 5.07 M (82.0% rooting). The number of ex vitro shoots formed was strongly correlated (r=0.66) with the length of roots formed in vitro, which was the highest at a 1.95 M ancymidol.     

Plant regeneration via somatic embryogenesis in Styrian pumpkin: cytological and biochemical investigations

Somatic embryo formation was induced from cotyledon explants of Styrian pumpkin (Cucurbita pepo L. subsp. pepo var. styriaca Greb.) by using a solid MS medium supplemented with 16.11M NAA and 4.44M BA or 26.85M NAA and 13.32M BA. The callus proliferation was more efficient on medium supplemented with 26.85M NAA and 13.32M BA. In contrast, the embryogenic response was higher on medium with lower concentrations of growth regulators (16.11M NAA and 4.44M BA). The time needed for embryo induction did not depend on medium composition. Embryos in globular stage were transferred to three different maturation media, containing 2.89M GA₃ in combination with 0.54M NAA, 11.42M IAA and growth regulator-free medium. The germination rate was the highest when embryos were cultured on medium with 11.42M IAA. Plantlets grown on this medium achieved maturity suitable for transplantation into soil within 9 to 10weeks. The regenerated plants were successfully transferred into field and developed fertile flowers and set fruits. Biochemical analysis showed significant lower total glutathione levels among in vitro grown plantlets compared to seedlings grown in soil. When the plantlets were transferred into soil, they reached a normal size within a month and the glutathione concentration was comparable to seed-derived plants at the same developmental stage. Transmission electron microscopy was used to investigate possible differences in the ultrastructure of cells from callus cultures, and leaf cells of regenerated and seed-derived plants. Differences in the ultrastructure were found within chloroplasts which contained only single thylakoids, large starch grains and small plastoglobuli in callus cells in comparison to leaf cells, which possessed a well developed thylakoid system, small starch grains and large plastoglobuli.     

In vitro plant regeneration via somatic embryogenesis from root culture of some rhizomatous irises

A method for plant regeneration of Iris via somatic embryogenesis is described. Root and leaf pieces from in vitro-grown plants of several genotypes of rhizomatous Iris sp. were cultured in vitro. Callus induction occurred only on root cultures incubated under low light intensity (35 mol m^-2 s^-1) on two induction media containing 2,4-D (4.5 or 22.5 M), NAA (5.4 M) and kinetin (0.5 M). Somatic embryos developed after transfer of callus onto four regeneration media containing 9 or 22 M BA, or 5 M kinetin and 2 M TIBA or 9 M BA and 4 M TIBA. Plantlets could be obtained from these somatic embryos. Genotypic differences were found both in callus induction and somatic embryo formation, with I. pseudacorus responding better than I. versicolor or I. setosa. Cytological analysis performed on root tips of 80 regenerated plants revealed that two of the I. pseudacorus regenerants were tetraploid.Abbreviations 2,4-D dichlorophenoxy acetic acid - NAA naphthaleneacetic acid - BA 6-benzyladenine - TIBA 2,3,5-triiodobenzoic acid - IBA indolebutyric acid     

The contribution of size-fractionated plankton to biomass and primary production of phytoplankton in saline–alkaline ponds

Primary productivity, biomass and chlorophyll-a of size fractionated phytoplankton (<0.22 m, <3 m, <8 m, <10 m, <40 m, <64 m, <112 m and <200 m) were estimated in 6 ponds and 5 experimental enclosures. The results showed that the planktonic algae less than 10 m are important in the biomass and production of phytoplankton in saline–alkaline ponds. The production of size fractionated phytoplankton corresponding to <112 m, <10 m and <3 m in saline–alkaline ponds were 10.5 ± 6.6 , 8.6 ± 5.4 and 0.33 ± 0.1 mgC l^–1 d^–1, respectively. Mean community respiration rate was 1.80 ± 0.73, 1.69 ± 0.90 and 1.38 ± 1.12 mgC l^–1 d^–1, respectively. The average production of phytoplankton corresponding to micro- (10–112 m), nano- (3–10 m) and pico- (<3 m) were 1.61, 8.30 and 0.33 mgC l^–1 d^–1, respectively. The ratio of those to the total phytoplankton production was 15%, 79% and 3%, respectively. The mean respiration rate of the different size groups was 0.11, 0.31 and 1.38 mgC l^–1 d^–1; the ratio of those to total respiration of phytoplankton was 6%, 17% and 77%, respectively. The production of size-fractionated phytoplankton corresponding to <200 m, <10 m and <3 m in enclosures was 2.19 ± 1.63, 2.08 ± 1.75 and 0.22 ± 0.08 mgC l^–1 d^-1, respectively. Mean community respiration rates were 1.25 ± 1.55, 1.17 ± 1.42 and 0.47 ± 0.32 mgC l^–1 d^–1, respectively. The average production of phytoplankton corresponding to micro- (10–200 m), nano- (3–10 m) and pico- (<3 m) plankton was 0.11, 1.86 and 0.22 mgC l^–1 d^–1, respectively. The ratio of those to the total production of phytoplankton was 5%, 85% and 10%, respectively. The mean respiration rate of different size groups were 0.08, 0.72 and 0.46 mgC l^–1 d^–1, the ratio of those to total respiration of phytoplankton was 6%, 57% and 37%, respectively. The concentrations of chlorophyll-a of the phytoplankton in the corresponding size of micro- (10–112 m), nano- (3–10 m) and pico- (<3 m) plankton in the experimental ponds were 19.3, 98.2 and 11. 9 g l^–1, respectively. The ratio of those to the total chlorophyll-a was 15%, 76% and 9%, respectively. The concentrations of chlorophyll-a of phytoplankton micro- (10–200 m), nano- (3–10 m) and pico- (<3 m) plankton in enclosures were 1.7, 34.3 and 3.0 g l^–1, respectively. The ratio of those to the total chlorophyll-a was 4%, 88% and 8%, respectively.     

Methylobacterium rhodesianum MB 126 possesses two acetoacetyl-CoA reductases

Two constitutive acetoacetyl-CoA (AcAc-CoA) reductases were purified from Methylobacterium rhodesianum MB 126, an NADPH-linked d(-)--hydroxybutyryl-CoA forming reductase (enzyme A) and an NADH-and NADPH-linked l(+)--hydroxybutyryl-CoA forming reductase (enzyme B). Enzyme A and B give apparent K _m values of 15 M and 30 M for AcAc-CoA, 18 M for NADPH and 30 M for NADH, respectively. They are inhibited by AcAc-CoA at concentrations higher than 25 M and 50 M, respectively. The contribution of the two reductases to poly--hydroxybutyrate synthesis is discussed.     

Theoretical considerations on growth kinetics and physiological adaptation of nutrient-limited phytoplankton

In many experimental ecological studies on phytoplankton species the growth response of an organism to the prevailing environmental conditions have been studied. Curves relating specific growth rate () to the external nutrient concentration (S) have been constructed to compare nutrient-limited growth of different species under steady state conditions. Microorganisms adapt their physiology to a certain limitation in order to optimize growth. Therefore the shape of the /S curve is closely related to the way a micoorganism adapts its physiology. To see how physiological adaptation and growth rate are interconnected to each other, both can be related to the internal concentration of the growth-limiting nutrient. How the growth rate is related to the internal nutrient concentration is presented in a mathematical equation.Many phytoplankton species during growth under different nutrient limitations show a linear relationship between and the reciprocal value of the internal nutrient content (= Yield). This was originally observed by Droop. The model presented here gives a theoretical explanation of this phenomenon.