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. 2026 Jun 17;208(7):81. doi: 10.1007/s00442-026-05922-5

The power of diet – the effect of different types of diet on the plastic body size response to temperature

Agata Burzawa 1,2,✉, Katarzyna Potera 1,2, Aleksandra Walczyńska 1
PMCID: PMC13275613  PMID: 42310231

Abstract

The temperature-size rule (TSR) describes the growth pattern of ectotherms, according to which they grow smaller at higher temperatures. While the general mechanisms of TSR are well studied, the role of diet in shaping these responses remains less understood. In this study, we examined how different diets influence the strength of TSR (= slope of the response) and the conditions under which it occurs. We used three clones of Lecane inermis rotifers of known thermal preferences: warm-preferring specialist, cold-preferring specialist and a generalist. The rotifers were cultivated at four temperatures, selected specifically for each clone, two lower, between which the rotifers switch from the negative to the positive growth rate (Tmin;8/10°C and 10/12°C), and two higher, around the expected optimal temperature at which fitness is the highest (Topt; 28 °C and 35 °C). The clone-specific thresholds were established in previous studies. The three laboratory diets were: a fluid with a known microorganismal content (Bio-Trakt bioproduct), a fluid that acts as a food source for bacterial biofilm (Molasses), and a powder that acts as a food source for bacterial biofilm (patented NOVO). We showed that the diets differently affected Tmin, Topt and the strength of TSR performance (measured as a comparison of body size within the range between Tmin and Topt). Of these three TSR response descriptors, Tmin was the most sensitive to diet, while Topt was the least. Our results show that different diets can affect the phenotypic plasticity patterns of organisms in different ways, and that different aspects of phenotypic plasticity (such as lower thermal limit, higher thermal limit, or strength of response) are sensitive to nutritional conditions in different ways.

Supplementary Information

The online version contains supplementary material available at 10.1007/s00442-026-05922-5.

Keywords: Body size, Diet, Ectotherms, Lecane inermis, Optimal thermal range, Phenotypic plasticity, Temperature-size rule (TSR)

Introduction

Temperature is a fundamental abiotic factor affecting all levels of biological organization (Schmidt-Nielsen, 1997). This influence begins at the level of the kinetics of biochemical reactions in living organisms (Hochachka and Somero, 2002). Through evolutionary processes, organisms have developed specific mechanisms to cope with diverse thermal environments (Somero 2010), including physiological, morphological, and behavioural mechanisms (Fangue et al. 2009). Importantly, organisms differ, sometimes considerably, in their ranges of thermal tolerance. This variability manifests at the molecular level, where some organisms are physiologically better adapted to particular temperatures than others (Fry, 1947). Following Levins (1968), such strategies are categorised as specialists, those that are better adapted to a specific range and perform worse in other conditions, and generalists, those that perform equally well over a relatively wide thermal range.

Ectotherms have evolved a number of physiological mechanisms to mitigate the possibly negative effects of temperature on their physiological processes and, consequently, their life histories (Angilletta 2009). One such response to changing temperature is to grow to larger sizes at lower (less preferable) temperatures, and to smaller sizes at warmer (more preferable) temperatures. This phenomenon is known as the Temperature-Size Rule (TSR), and was observed in more than 80% of ectothermic organisms, covering both terrestrial and aquatic species (Atkinson 1994; Table 1).

Table 1.

The taxonomic distribution and environmental context of TSR studies

Organisms Genres Environment References
Fish haddocks, cods, Norway pouts marine Baudron et al. 2014; Coghlan et al. 2024; Shapiro Goldberg et al. 2019; van Rijn et al. 2017
Amphibians salamanders, anurans freshwater Rollinson and Rowe 2018; Ruthsatz et al. 2018; Zhu et al. 2023
Crustaceans planktonic crustaceans, Daphnia spp. freshwater Jalal et al. 2013
Insects flies, bees, Drosophila melanogaster, Osmia bicornis terrestrial Blanckenhorn and Llaurens 2005; Czarnoleski et al. 2013; Ghosh et al. 2013; Kierat et al. 2017
Nematodes Caenorhabditis elegans terrestrial Kammenga et al. 2007; Van Voorhies 1996
Reptiles lizards, turtles terrestrial, marine Aragón and Fitze 2014; Czarnoleski et al. 2017; Santilli and Rollinson, 2018
Rotifers rotifers freshwater Czarnoleski et al. 2015; Kiełbasa et al. 2014; Stelzer 2002; Walczyńska et al. 2015
Diatomes phytoplankton freshwater, marine Peter and Sommer 2012; Walczyńska and Sobczyk 2017; Winder et al. 2009
Protists protists freshwater Atkinson et al. 2003; Walczyńska et al. 2016
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The TSR is a “rule” and not a “law” as many contradictory examples have been observed (e.g., Atkinson 1995; Walters and Hassall 2006). Such findings call into question the universality of the TSR and the existence of a general mechanism driving such growth patterns (Angilletta and Dunham 2003). However, the explanation may be much simpler than we expect. According to the recent studies on rotifers (Walczyńska et al. 2016; Żygadło et al. 2025), which have been confirmed for the case of insects (Blanckenhorn et al. 2021), the TSR occurs only under certain conditions, enclosed between the minimal and optimal temperature for population growth rate, Tmin and Topt, respectively (Walczyńska et al. 2016; Żygadło et al. 2025). Beyond this range, body size exhibits similar values at both thermal ends (see Fig. 3 in Walczyńska et al. 2016) and lies around a zero-plasticity line (see Fig. 4 in Walczyńska et al. 2016). Such a pattern suggests that body size beyond a certain optimal thermal range is canalised sensu Waddington (1942). From an evolutionary perspective, the canalisation of a trait ensures a stable reaction norm across environments (Stearns et al. 1995; Stearns and Kawecki, 1994). The switches from canalisation to plasticity that occur at different temperatures cause the pattern around Tmin and Topt that, at first sight, appears to be the opposite of TSR (Fig. 1A). However, this arises from the existence of energetic costs of phenotypic plasticity. Recently, such costs were detected at the level of cellular respiration as associated with the increasing costs of mitochondrial performance and non-mitochondrial respiration at higher temperature (Burzawa et al. 2025). With this knowledge, at least some of the observed deviations from the TSR can be explained by the fact that the thermal preferences of the organisms were not taken into account in the selection of thermal conditions during the study (Walczyńska et al. 2016).

Fig. 3.

Fig. 3

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The mean values of body size for each treatment over temperature. The dots in the colours of the respective diets represent the individual body size measurements, boxes denote interquartile range and whiskers represent the standard error of each temperature and treatment. The numbers above the boxes correspond to the sample size. Body size was estimated as the product of width and length as a proxy of rotifer area [µm2]

Fig. 4.

Fig. 4

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Effect size analysis using Cohen’s d with 95% Confidence Intervals. Accepted interpretation of Cohen’s d values (Cohen J., 1988) was |0.20| ≥ d ≤ |0.50|: small effect; |0.51| ≥ d ≤ |0.80|: medium effect; d ≥ |0.81|: large effect. The effect size is significant when CIs do not include 0. N.S. – not significant

Fig. 1.

Fig. 1

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Schematic representation of the hypotheses tested in the study. Panel A shows a body size pattern for within and beyond the optimal thermal range for the TSR, enclosed between Tmin and Topt for fitness, as established in the previous studies (Walczyńska et al. 2016; Żygadło et al. 2025). The red dots show the sizes expected at selected temperatures. Panel B shows possible changes in the strength of the TSR expected if diet has an effect, as predicted in hypothesis (i). Panels C and D show possible shifts in Tmin and Topt, respectively, if affected by different diets, according to hypothesis (ii)

The optimal conditions for the occurrence of TSR are organism-specific and potentially relate to all parameters to which an organism is exposed in its environment, not just temperature (Kiełbasa et al. 2014; Kutz et al. 2019). Another factor that makes the observed relationships more difficult to understand is the interdependence between environmental parameters. This is not surprising, as ectotherms only behave alike at a given temperature if all other environmental parameters are held constant (Sinclair et al., 2016). Referring to the effect on the TSR pattern, the most important environmental parameters besides temperature are nutrient conditions and oxygen availability; depending on the strength of the relationship between temperature and either nutritional conditions or oxygen availability, the TSR pattern may or may not be observed (Walczyńska and Serra 2022; Walczyńska and Sobczyk 2017). The quality of the diet is particularly important in this respect (Bazin et al. 2023; Berrigan and Charnov 1994; Diamond and Kingsolver 2010; Kiełbasa et al. 2014; Kutz et al. 2019). In the field, many ectotherms shift their diets as the ambient temperature changes, presumably because the optimal protein/carbohydrate ratio is temperature-dependent (Hardison et al. 2021). This feature is of great ecological importance because, according to the modern stoichiometric approach in ecology, changing the ratio of protein to carbohydrate in a diet leads to changes in multi-element stoichiometry (El-Sabaawi et al. 2023; Filipiak and Filipiak 2022; Peñuelas et al. 2019). Another possible explanation for the potential effect of diet on plastic body size adjustment to temperature is that the quality of different types of diet may change differently with temperature, depending on whether the diet consists of organisms with their own specific thermal preferences (e.g., Eskuche-Keith et al. 2024), or of substances that may act as a biofilm for uncontrolled communities of microorganisms (Horváthová et al., 2016, 2019; Horváthová and Bauchinger 2019). Such effects have not been rigorously tested, but the first step is to find out if and how different types of standard laboratory diets can indeed affect the pattern of plastic response of body size to changing temperature. This preliminary question is rigorously addressed in this study.

Although the individual effects of temperature and diet on different organismal traits have been well studied, their combined effects have only recently been explored, as reviewed by Hardison and Eliason (2024). In the context of the TSR, poor quality diet was found to affect or even reverse the growth pattern in the rotifers Lecane inermis (Kiełbasa et al. 2014) and Euchlanis dilatata (Santos-Medrano et al. 2022), the bowfly Aldrichina grahami (Yan et al. 2024), the butterfly Manduca sexta (Diamond and Kingsolver 2010), and the amphibian Rana temporaria (Liess et al. 2013). Additionally, the TSR was found to be influenced by the macronutrient balance of a diet in Drosophila melanogaster (Kutz et al. 2019) and the butterfly Spodoptera litura (Lee et al. 2015).

Our aim was to investigate whether the general pattern of the organism-specific TSR is resistant to dietary conditions. We used rotifer Lecane inermis, the species that follows the TSR not only under controlled laboratory conditions but also in field (Kiełbasa et al. 2014). For the case of L. inermis, the TSR was found to be an adaptation to temperature-dependent oxygen conditions (Walczyńska et al. 2015) realized only within a certain thermal range (Walczyńska et al. 2016; Fig. 1). We fed rotifers three types of standard laboratory diets and we posed the following questions: (i) does diet affect the strength of the TSR (Fig. 1B)? (ii) does diet shift the temperature thresholds within which the TSR occurs (Fig. 1C and D)? In addition, population growth rate was included as a proxy for fitness to enable evolutionary interpretation of the observed patterns.

Understanding how food may potentially affect the plastic body size response to temperature changes is relevant in the context of climate change, as shifts in temperature regimes may have implications for fitness of ectotherms, potentially affecting their ecological roles and interactions within ecosystems.

Methods

Culture maintenance

The model organism in this study was Lecane inermis Bryce (Rotifera, Monogononta). This bacterivorous species is commonly found in freshwater environments and plays a crucial role in regulating microorganism populations as well as serves as prey for larger aquatic organisms, highlighting its importance in maintaining ecological equilibrium in freshwater ecosystems (Fiałkowska et al. 2011). The clones used in this experiment were subjected to a prior assessment of their TSR performance over a wide thermal spectrum, and their thermal preferences were estimated (Stuczyńska et al. 2022; Żygadło et al. 2025). From among those clones, we used three which differed regarding their known thermal preferences; one showing a specialization towards lower temperatures, herein referred to as ‘COLD’, one showing specialization towards high temperature, herein referred to as ‘WARM1’, and the third representing a generalist clone, characterized by relatively good performance across the broad thermal range, here denoted as ‘INT’.

Experimental conditions

The temperature regimes used in the study were selected on the basis of previously established clone-specific thresholds for the minimum temperature for proliferation, Tmin, and the optimum temperature at which the clone proliferates best, Topt (Table 3 in Żygadło et al. 2025). According to the results of that study, the temperatures around Tmin were set at 8 °C and 10 °C for the COLD and INT clones, and at 10 °C and 12 °C for the WARM1 clone. In the case of Topt, all clones were exposed to 28 °C and 35 °C (Fig. 2).

Fig. 2.

Fig. 2

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Schematic diagram showing the order in which the procedures were carried out; which clone (COLD, INT or WARM1; first row) was examined in which thermal conditions (cold or warm; second row) and how it relates to the timeline (in days; third row). Day 0 refers to the starting point of the experiment. Fitness measurement I and II refer to two counts of rotifers that were used to estimate fitness (description in the text). M - Molasses, B – Bio-Trakt, N – NOVO

We used three types of diet, each of which is used in the daily maintenance of our lab culture, but which differ in form and composition:

  1. NOVO® is a powder formulation serving as a food for rotifers, patented as an optimally balanced food source (Pajdak-Stós et al. 2016). In our laboratory it is a predominant type of diet used to maintain rotifer cultures (hereafter, diet N).

  2. Molasses (3‰) was used as the standard liquid supplement commonly applied in zooplankton culture (Abedini Najafabadi et al. 2015; Pajdak-Stós et al. 2011). In our setup, molasses provided an extra source of organic carbon that was easily accessible to heterotrophic bacteria. This promoted the growth of heterotrophic bacteria, which then formed the basis of a microbial food source for the rotifers (hereafter, diet M).

  3. Bio-Trakt® (Zielone Oczyszczalnie, Poland) is a commercially available liquid bioproduct designed to support biological decomposition processes in aquatic systems. Unlike the other two diets, it contains a known microbial composition, including both prebiotic and probiotic components: lactic fermentation bacteria, radiata and phototrophic bacteria, yeast, ethanol, natural acetic acid, organic sugar cane molasses, pure non-chlorinated water (hereafter, diet B).

Rotifers were maintained in 60 mm ⌀ Petri dishes in 10 ml of Żywiec Zdrój® spring water (Poland). We started the experiment with 20 individuals per dish in three replicates per treatment. The amount of food in each dietary regime was determined in a pilot study to ensure ad libitum availability, allowing rotifers to feed without restriction. The rotifers were then fed 300 µl of molasses, 50 µl of Bio-Tract® or one pinch (approximately 3 mg) of NOVO® on day 0 of the experiment and again when the experimental populations were distributed across different temperatures (day 3 in Fig. 2). Thus, acclimation to the experimental conditions occurred in two stages; from the experiment onset on day 0 until the third day, the rotifers were acclimated to the experimental diet, while between the third and sixth day, they were subsequently acclimated to the target temperature (Fig. 2). For the logistical reasons, each clone was investigated separately.

Measurements

Two traits were examined, body size and population growth rate r. Body size served as an indicator of compliance with the TSR, while r was used as a proxy for fitness, thus providing insight into the adaptive responses of organisms under the given experimental conditions. We also needed fitness to control the thermal thresholds in order to assess whether the effect was in accordance with the ‘optimal thermal range’ assumptions (see Fig. 1).

Fitness was calculated based on two counts of rotifers in each diet and temperature combination. Population growth rate measurements began on day 6. In the warm-temperature treatments, growth rate was assessed on days 6 and 8, whereas in the cold-temperature treatments it was assessed on days 9 and 13. Fixation with Lugol’s solution was performed on day 9 for the warm-temperature treatments and on day 14 for the cold-temperature treatments. For higher temperatures, five 25-µl subsamples were taken from each well as a standard method (Dubber and Gray, 2009). The rotifers were counted using the light microscopy. The mean rotifer density per sample was calculated from the three values remaining after removal of the highest and the lowest value. For the lower temperatures, the rotifers were counted directly from the wells (all rotifers per well).

After the first and second counts animals were fed ad libitum according to their diet regime (third and fourth feeding). The population growth rate r was calculated as the difference between the number of rotifers from the second and the first counts, divided by the period of time between counts in days:

graphic file with name d33e821.gif

where: r – population growth rate, n1 – number of rotifers in first count, n2 – number of rotifers in second count, t – time period between two counts [days].

The pattern obtained for r serves as a test of whether the conditions in the experimental cultures were optimal for the rotifers’ growth. If so, we expect the fitness to increase with increasing temperature until the optimum Topt is reached and then to decrease at the highest temperature.

After the second count, the rotifers were fed again and fixed with Lugol’s solution the next day. For body size measurements, approximately 50 individuals were selected from each temperature and diet combination, pooled across replicates. Body size was estimated as the product of width and length as a proxy of rotifer area (Fig. 2).

Statistical analyses

Each clone was analyzed separately in R programming language (R Core Team, 2020). In all the analyses, the natural logarithm of body size was used. To address whether diet affects the strength of the TSR response (hypothesis (i)), we compared body size for two temperatures assumed to be within a thermal range for the TSR, as presented in Fig. 1B. Thus, for the COLD and INT clones, body size was compared for 10 °C and 28 °C, and for the WARM1 clone for 12 °C and 28 °C. Previous studies have estimated the strength of the TSR as the slope of the linear response within the optimal thermal range (Żygadło et al. 2025). In this study, where two body sizes are being compared, the strength is represented by the difference (effect size) between them.

To test whether diet shifts the lower (Tmin) or upper (Topt) thermal thresholds for the TSR (hypothesis (ii)), we compared body size around lower thermal threshold at 8 °C and 10 °C for the INT and COLD clones, and at 10 °C and 12 °C for the WARM1 clone (Fig. 1C and D, respectively). Referring to the upper thermal threshold, we compared body size and r at 28 °C and 35 °C.

Because statistical significance alone does not fully capture the biological relevance of the results, we additionally estimated effect sizes to evaluate the magnitude and consistency of the treatment responses. The effect size analyses were estimated using effect size packages (Ben-Shachar et al. 2020). For this purpose, we calculated Cohen’s d to quantify the magnitude of the differences in body size between pairs of temperatures for the three diets.

We followed the instruction for accepted interpretations of Cohen’s d values according to Cohen (1988):

|0.20| ≥ d ≤ |0.50| : small effect.

|0.51| ≥ d ≤ |0.80|: medium effect.

d ≥ |0.81|: large effect.

The upper and lower CI values inform about statistical significance of a given effect. The effect size is significant when CIs do not include 0.

Results

The results demonstrate the differential effects of the three standard diets used in the mass culture of the examined species on the determinants of the plastic body size response to temperature. These determinants are the strength of the response and the low and high thermal thresholds at which the response occurs (presented schematically in Fig. 1B, C and D, respectively). The TSR trait that was the most strongly affected was the minimum response temperature, followed by response strength. The upper limit of the response was least affected by the diet (Fig. 4).

Descriptive results

The dataset of body size was subjected to outlier detection using boxplot analysis (ggplot2 packages, Wickham, 2016) and any identified outliers were excluded from further analyses. Due to a handling error, one replicate of the INT clone at 35 °C on diet B was discarded from the analyses. Body size measurements for the INT clone around Tmin fed diet N were not possible due to poor rotifer proliferation. The average body size per clone, temperature, and diet is shown in Supplementary Table S1, while its distribution is presented in Fig. 3. The TSR pattern predicted by the ideal scheme shown in Fig. 1A can be seen in some, but not all, cases. The clearest pattern across clones is for WARM1, while across diets it is for diet B (see Fig. 3).

Effect size analysis

Effect size analysis quantifies the magnitude of a biological response, providing information that goes beyond statistical significance alone (Spake et al. 2023). In ecological and evolutionary studies—where variability is high and sample sizes often differ—effect sizes allow meaningful comparison across treatments. To test whether diet affected the strength of the TSR, we compared body size within thermal thresholds. The pattern of TSR should show a negative Cohen’s d, as within the thermal threshold, body size should decrease with increasing temperature (Fig. 1). Such a pattern was found for diet B for all three clones and for diets M and N for clone WARM1, with no differences between diets (Fig. 4).

To investigate the possible effect of diet on the lower thermal threshold for TSR, we compared body size for two lowest temperatures. If the threshold was correctly captured, Cohen’s d should be positive, as body size below the threshold should decrease to the canalized value (Fig. 1). The positive Cohen’s d would mean that the actual thermal threshold was at lower temperature (Żygadło et al. 2025). We found the positive Cohen’s d for diet B in clones INT and COLD and for diets M and N in clone WARM1. For diet B in clone WARM1 the effect was negative. All other effects were not significant (Fig. 4).

For the effect of diet on the higher threshold for TSR, we compared body size for the two higher temperatures and expected a positive Cohen’s d (Fig. 1). In all cases the effect was not significant, except in the case of diet M in clone COLD where the effect was negative (Fig. 4). All Cohen’s d values and their CI are provided in supplementary Table S2.

Reference to fitness

The results for r as a measure of fitness show that the expected pattern was generally maintained, with the most notable exceptions for the M and N diets in the COLD clone, where r at 10 °C was clearly lower than at 8 °C. Such a result may suggest the presence of an interfering, uncontrolled factor that may affect the ability to respond plastically. Therefore, the body size of rotifers in these treatments should be treated with caution. At lower temperatures, all diets provided similar fitness, while diet N resulted in considerably higher fitness than the other two diets at higher temperatures (Fig. 5).

Fig. 5.

Fig. 5

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The mean values of population growth rate r as a proxy of fitness for each treatment over temperature. The sample size in each temperature treatment was 3. Whiskers represent the standard error

Discussion

We show that even a standard laboratory diet can influence the conditions under which phenotypic plasticity is realized. In the case of our study, phenotypic plasticity was the ability of an organism to plastically change its body size in response to changing temperature, while the diet represented three variants differing in consistency and composition, all used in laboratory cultures and resulting in positive rotifer proliferation.

We tested the difference between the diets in the body size pattern at a lower limiting temperature for the TSR performance (the thermal threshold around which the rotifers were expected to switch from negative to positive population growth rate; Tmin), a higher limiting temperature (the threshold above which rotifers were expected to have lower fitness; Topt) and the strength of the plastic response within the optimal thermal range (Fig. 1). We found that Tmin was the most sensitive to the dietary conditions, with large and significant (positive change) effect of the Bio-Trakt diet on the body size of the INT and COLD clones and medium or large effect of all three diets on the body size of the Warm1 clone (negative change in the Bio-Trakt and positive change in the other two diet regimes) (Fig. 4). Regarding the strength of the TSR, the Bio-Trakt diet had a greater effect on body size differences (stronger response) in the INT and COLD clones than other diets did. A large negative effect on TSR (stronger response) was observed in the WARM1 clone, regardless of the dietary regime (Fig. 4). Topt was the most resistant to different diets, with only one significant (medium) effect on the body size of the COLD clone in Molasses (Fig. 4).

The most pronounced effect was observed in the Bio-Trakt diet, as TSR patterns were most evident in this diet. These patterns showed a decrease in body size with increasing temperature within the range that defines the TSR pattern, and a canalisation of body size outside this range, as predicted (Fig. 1A). Secondly, Bio-Trakt clearly shifted the minimum temperature for the WARM1 clone, as evidenced by the reversed body size pattern observed at two lower temperatures (Fig. 3). This suggests the presence of phenotypic plasticity at a lower temperature than that set in the experiment. Comparing the other two diets, no differences were found in the effects of Molasses or NOVO, except in one case; NOVO caused significantly larger effect on the pattern reverse to TSR around Tmin in WARM1 (Figs. 3 and 4). This suggests that the temperatures chosen for Tmin were best set for the NOVO diet for this clone. The effect of Molasses was mostly intermediate among the diets.

An interesting result is that the diet that resulted in the highest fitness of the three diets, NOVO, at least around optimal temperature, was not the one that most affected phenotypic plasticity. The distinguishing feature of the most influential diet, Bio-Trakt, was that it contained microbial populations, while the other two diets were a composition of nutrients of various types acting as a food source for rotifers and/or bacteria as a food for rotifers. It may be the reason why this diet caused the clearest plastic pattern, especially at low temperature. It appears that the consistency of the diet, whether powdered, as with NOVO, or liquid, as with the other two diets, had no effect on phenotypic plasticity.

The physiological limits of thermal tolerance, usually expressed as critical values CTmin and CTmax, are generally considered as constant (Anthony et al. 2021; Morgan et al. 2018). However, we demonstrate that diet can modulate the thermal thresholds expressed as ecological measures, Tmin and Topt. This indicates that thermal responses are not solely determined by physiological boundaries, but are also driven by ecological limits that arise from resource-dependent physiological performance. This refers specifically to Tmin, and this result in consistent with the results of Bazin et al. (2023), who found that food restriction had more severe effect at lower than at higher temperature in medaka fish tested for the joint effect of temperature and diet on the TSR pattern. Furthermore, this is consistent with the more general pattern observed in ectotherms, which demonstrate greater variability in their tolerance to cold than to heat. This suggests that tolerance to cold is more flexible and susceptible to natural selection than the physiologically conserved tolerance to heat (Araújo et al. 2013; Walczyńska et al. 2021).

Our results are important in the context of laboratory studies, where diets are typically standardized and artificial. We demonstrate that the phenotypic plasticity of organisms may vary depending on the nutritional content of their food, even when the diets used in experiments are optimal and result in sufficient proliferation. Particular attention should be paid to diets containing living organisms, as these can have different thermal preferences to their consumers, introducing an uncontrolled interfering factor. Even more importantly, variation in thermal limits of organisms at different levels of a trophic food web of the natural community may cause considerable and unpredictable changes to community structure under conditions of rapid temperature changes (Katzenberger et al. 2021).

Diet was found a determinant of the thermal performance of ectotherms in different traits and in different way (Hardison et al. 2021; Kutz et al. 2019). To the best of our knowledge, the combined effect of these two factors has not yet been tested for the limits of phenotypic plasticity. Since the plastic response is the organismal first line of defense against environmental changes (e.g., Levis and Pfennig 2016), our study is an important contribution to understanding of the mechanisms that limit the organismal ability to respond to changing conditions. In addition, we relate the tested patterns to the population growth rate as a direct measure of fitness, which makes it possible to interpret the results in an evolutionary context.

The fitness generally increased with temperature up to Topt and then decreased, confirming that experimental conditions enabled the optimal growth of rotifers. The visible deviation from those pattern is for COLD clone kept at 10 °C and fed Molasses and NOVO. It may have affected the body size of rotifers in that treatments and the respective body size results should be treated with caution, however, the experimental cultures did not show the signs of bad conditions. Another reason for such a result could be that population growth rate estimation is less precise at lower than at higher temperatures because of low rotifer densities.

To sum up the contribution of our results, the importance of both temperature and diet as key factors affecting the physiology and life-history traits of ectotherms is increasingly recognized in the context of the so-called nutritional or digestive physiology (Hardison and Eliason 2024 and citations therein). This is particularly important in natural communities exposed to profound changes caused by humans in a variety of ways (Birnie-Gauvin et al. 2017).

Supplementary Information

Below is the link to the electronic supplementary material.

Supplementary Material 1 (25.4KB, docx)

Acknowledgements

Authors thank Agnieszka Pajdak-Stós and Edyta Fiałkowska for their kind support and expert guidance in rotifer cultivation and for valuable comments on the previous version of the text.

Authors contribution

AW conceived the ideas and designed methodology and revising it critically for important intellectual content; AB conceived the ideas and designed methodology, collected the data, analysed the data and led the writing of the manuscript; KP collected the data. All authors contributed critically to the drafts and gave final approval for publication.

Funding

This work was supported by the National Science Centre of Poland (2019/35/B/NZ8/ 03245).

Data availability

The data and code that support the findings and analyses of this study are available at https://uj.rodbuk.pl/dataset.xhtml?persistentId=doi:10.57903/UJ/MP9O0G.

Code availability

The code that support the findings of this study will be made publicly available in an open-access repository upon acceptance and publication of the article in open repository of Jagiellonian University.

Declarations

Conflicts of interest

The authors declare that they have no conflict of interest.

Ethics approval

For this type of study formal consent is not required.

Consent to participate

Not applicable.

Consent for publication

Not applicable.

Footnotes

Publisher’s Note

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Associated Data

This section collects any data citations, data availability statements, or supplementary materials included in this article.

Supplementary Materials

Supplementary Material 1 (25.4KB, docx)

Data Availability Statement

The data and code that support the findings and analyses of this study are available at https://uj.rodbuk.pl/dataset.xhtml?persistentId=doi:10.57903/UJ/MP9O0G.

The code that support the findings of this study will be made publicly available in an open-access repository upon acceptance and publication of the article in open repository of Jagiellonian University.

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