How does competition shape variation in metabolic rates?

A new study pub­lished in Evol­u­tion Let­ters invest­ig­ates why meta­bol­ic rates are so vari­able between organ­isms. Here, lead author Dr Aman­da Pet­tersen tell us what she found.

Meta­bol­ic rate – the rate at which organ­isms take up, trans­form, and expend energy, is linked with the “pace-of-life”. Indi­vidu­als with high meta­bol­ic rates often grow faster, repro­duce soon­er, and are short­er lived, com­pared with indi­vidu­als with low meta­bol­ism that are often longer lived. Dif­fer­ences in energy expendit­ure among indi­vidu­als are likely to res­ult in dif­fer­ences in sur­viv­al and repro­duc­tion, so we expect meta­bol­ic rates are under selec­tion. Evol­u­tion­ary the­ory pre­dicts that over time, selec­tion should deplete vari­ation in traits, yet even after account­ing for body size and tem­per­at­ure, we can observe a threefold dif­fer­ence in meta­bol­ic rates across indi­vidu­als of the same spe­cies. How can we account for this variation?

The pres­ence and mag­nitude of biot­ic and abi­ot­ic factors can vary con­sid­er­ably in nat­ur­al envir­on­ments, even across small spa­tial scales. Dif­fer­ences in selec­tion regimes across envir­on­ments may main­tain vari­ation in meta­bol­ic rates, even under strong selec­tion in any single envir­on­ment. This is because a fast or slow pace-of-life might be advant­age­ous depend­ing on the envir­on­ment an organ­ism inhab­its. While vari­ation in selec­tion over large and small spa­tial scales have been found pre­vi­ously, field estim­ates of selec­tion on meta­bol­ism across envir­on­ments are his­tor­ic­ally rare.

The colo­ni­al bryo­zo­an, Bug­ula ner­it­ina. Image: Aman­da Pettersen.

To invest­ig­ate the role of envir­on­ment­al vari­ation in main­tain­ing trait vari­ation, we stud­ied the mod­el mar­ine inver­teb­rate, Bug­ula ner­it­ina. The adult stage of Bug­ula ner­it­ina is a colo­ni­al bryo­zo­an, com­prised of thou­sands of indi­vidu­al sub­units (zooids). Some zooids con­tain repro­duct­ive struc­tures (ovi­cells) that hold indi­vidu­al lar­vae. In response to light expos­ure at dawn, the par­ent colony releases hun­dreds of swim­ming lar­vae into the plank­ton – the lar­vae soon find a suit­able sub­strate to settle on, such as a pier pylon, and meta­morph­ose into the first zooid of the colony. From here, the zooids begin bud­ding and the colony grows, form­ing ovi­cells approx­im­ately four weeks after settlement.

We settled indi­vidu­al mar­ine bryo­zo­an lar­vae by pipet­ting them in a drop of water onto small pieces of acet­ate (as a set­tle­ment sub­strate) and meas­ured their meta­bol­ic rates. We meas­ured meta­bol­ic rate (rate of oxy­gen con­sump­tion) at two time points: 2 hours (“Meta­bol­ic Rate Early”; MRE) and 24 hours (“Meta­bol­ic Rate Late”; MRL) post-set­tle­ment. We then glued the set­tlers on the acet­ate pieces onto tiles in one of three com­pet­i­tion envir­on­ments: no com­pet­i­tion (“nocomp”), with­in-spe­cies com­pet­it­ors (“intra”), and a pre-estab­lished mar­ine com­munity (“inter”). We ran­domly assor­ted these treat­ment tiles onto 10 back­ing pan­els (36 tiles per pan­el) and hung them at 1.5m depth in the sub­tid­al at our field site – a mar­ina in Mel­bourne, Aus­tralia. We mon­itored indi­vidu­als approx­im­ately weekly for 4 months, meas­ur­ing fit­ness: sur­viv­al (“viab­il­ity”), the abil­ity to repro­duce (“fer­til­ity”), and repro­duct­ive out­put (“fecund­ity”), along with our pace-of-life traits: growth, age at onset of repro­duc­tion, and longevity.

We found that selec­tion on meta­bol­ic rates (MRE and MRL) var­ies among com­pet­i­tion envir­on­ments sep­ar­ated by only a few cen­ti­metres – com­pet­i­tion selects for a faster pace-of-life, com­pared with com­pet­i­tion-free envir­on­ments. High-meta­bol­ism indi­vidu­als are bet­ter able to with­stand intense com­pet­i­tion from both with­in-spe­cies com­pet­it­ors and in a pre-estab­lished mar­ine com­munity. How­ever, low-meta­bol­ism indi­vidu­als live longer, and showed high­er fit­ness under com­pet­i­tion-free con­di­tions. If arriv­ing into a com­pet­i­tion-free space, indi­vidu­als will have high­er fit­ness if they grow slow and repro­duce later but live longer. Hence, the envir­on­ment-depend­ent nature of selec­tion on meta­bol­ism and the pace-of-life is likely to main­tain vari­ation in meta­bol­ic rates.

These find­ings sup­port pre­vi­ous ground­work in field eco­logy, sug­gest­ing that dif­fer­ences in selec­tion regimes main­tain trait vari­ation. We found that selec­tion on meta­bol­ic rates can vary con­sid­er­ably, even over a few cen­ti­metres! A high meta­bol­ic rate and fast pace-of-life is favoured under strong com­pet­i­tion – indi­vidu­als die young but repro­duce soon­er. How­ever, in the rare case that indi­vidu­als find them­selves in a com­pet­i­tion-free envir­on­ment, then a low meta­bol­ic rate and slow pace-of-life can provide a fit­ness advantage.

Dr Aman­da Pet­tersen is a postdoc­tor­al research­er at the Uni­ver­sity of Lund. The ori­gin­al art­icle is freely avail­able to read and down­load from Evol­u­tion Let­ters.

You can also watch a short talk by Dr Pet­tersen on the study here: