The growth rate hypothesis (GRH) posits an increase in ribosomal ribonucleic acid (RNA) content, and therefore cellular phosphorus (P), with increasing growth rate. There is evidence that the GRH may not apply to phytoplankton under all conditions. Here, we experimentally controlled four conditions (light, temperature, pH, and CO2) to alter the growth rate of Emiliania huxleyi, a biogeochemically important coccolithophorid, and monitored changes in RNA, protein, and carbohydrate content. We show that an increase in growth rate caused by increasing light, pH, and CO2 resulted in increased RNA per unit of organic carbon (RNA : POC), but that increasing temperature, leading to increase of growth rate, resulted in a decrease in RNA : POC. Protein per unit of organic carbon (protein : POC) increased in our increased temperature, pH, and CO2 treatments that increased growth rate, but there was little change in protein : POC in our light treatment despite it inducing the same increase in growth rate. Carbohydrate per unit of organic carbon (Carbohydrate : POC) increased with growth rate under increased light and CO2 but did not vary significantly in the temperature or pH treatments. These results indicate that physiological acclimation to specific environmental conditions can lead to contrasting patterns in RNA, protein, and carbohydrate composition and therefore contrasting changes in carbon : nitrogen : phosphorus ratios with growth rate in E. huxleyi.

Housekeeping genes (HKGs) are constitutively expressed with low variation across tissues/conditions. They are thought to be highly conserved and fundamental to cellular maintenance, with distinctive genomic features.Here, we identify 1505 HKGs in the unicellular marine diatom Thalassiosira pseudonana based on an RNA-seq analysis of 232 samples taken under 12 experimental conditions over 0-72 h. We identify promising internal reference genes (IRGs) for T. pseudonana from the most stably expressed HKGs.A comparative analysis indicates < 18% of HKGs in T. pseudonana have orthologs in other eukaryotes, including other diatom species. Contrary to work on human tissues, T. pseudonana HKGs are longer than non-HKGs, due to elongated introns. More ancient HKGs tend to be shorter than more recent HKGs, and expression levels of HKGs decrease more rapidly with gene length relative to non-HKGs.Our results indicate that HKGs are highly variable across the tree of life and thus unlikely to be universally fundamental for cellular maintenance. We hypothesize that the distinct genomic features of HKGs of T. pseudonana may be a consequence of selection pressures associated with high expression and low variance across conditions.

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