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It Pays to Be Different:
Evolutionary Distinctiveness and Conservation Priorities

By Claire Asher, on 15 July 2014

The world is currently experiencing an extinction crisis. A mass extinction on a scale not seen since the dinosaurs. While conservationists work tirelessly to try and protect the World’s biodiversity, it will not be possible to save everything, and it is important to focus conservation efforts intelligently. Evolutionary distinctiveness is a measure of how isolated a species is on it’s family tree – how long ago it split off from its nearest living relative. A recent paper co-authored by UCL GEE’s Dr David Redding, published in Current Biology, assessed how effective evolutionary distinctiveness is a tool for identifying bird species of conservation priority. Current conservation efforts are missing some of the most evolutionarily distinct species.

Evolutionary distinctiveness (ED) is measured as the distance along the evolutionary tree from one species to it’s nearest relative. It can be used as a measure of how much evolutionary ‘information’ would be lost if this species were to become extinct. We have good estimates of these distances for birds as we have been able to put dates on the evolutionary tree based on fossil records and molecular data. A recent analysis of nearly 10,000 known bird species, by researchers at Yale University, Imperial College London, University of Sheffield, Simon Fraser University, University of Tasmania and University College London, showed some patterns we might have expected, for example, evolutionary distinctiveness is highest in isolated regions (e.g. Australia, New Zealand and Madagascar) and regions with higher species richness tended to have more evolutionary distinct birds. However, there were also some unexpected results. For example, ED wasn’t strongly related to latitude, a pattern predicted by the idea that the tropics act as a ‘museum’ for ancient lineages, nor was ED related to a species’ range-size, which has previously been predicted theoretically.

Evolutionary distinctiveness showed little relationship with conservation status – some of the most threatened distinct species are found outside of biodiverse regions that are usually the target of conservation efforts. This means that, when we consider only species richness or total biodiversity to identify regions to conserve, we may be missing a great deal of evolutionary information. Instead, basing areas of conservation priority on the evolutionary distinctiveness of their flora and fauna may offer a more efficient and effective way to maximise the evolutionary variation we keep.

The paper also released the first formal list of ‘EDGE birds’ – EDGE stands for “Evolutionary Distinctive and Globally Endangered” and is a metric combining ED with the IUCN Red List. The list includes the Giant Ibis, the New Caledonian Owlet-Nightjar, the California Condor, the Kakapo, the Philippine Eagle, the Christmas Island Frigatebird and the Kagu, all of which are listed as either Critically Endangered or Endangered.

The most evolutionary distinct birds include both common species and rare species, both isolated and wildly distributed species, and are found in almost every environment on Earth. Current conservation efforts that focus on tropical regions with high species richness may be neglecting many evolutionary distinct species, whose extinction would represent the loss of a great deal of ‘evolutionary information’. Evolutionary distinctiveness could offer a powerful tool to supplement current criteria for identifying conservation priorities.

Original Article:

() Current Biology

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This research was made possible by funding from the Natural Environment Research Council (NERC), the Natural Sciences and Engineering Research Council (NSERC), the National Science Foundation, and the National Aeronautics and Space Administration (NASA)

Synthetic Biology and Conservation

By Claire Asher, on 7 July 2014

Synthetic biology, a hybrid between Engineering and Biology, is an emerging field of research promising to change the way we think about manufacturing, medicine, food production, and even conservation and sustainability. Oryx front cover
A review paper released this month in Oryx, authored by Dr Kent Redford, Professor William Adams, Dr Rob Carlson, Bertina Ceccarelli and CBER’s Professor Georgina Mace, discusses the possibilities and consequences of synthetic biology for biodiversity conservation. Synthetic biology aims to engineer the natural world to generate novel parts and systems that can be used to tackle real world problems such as genetic disease, food security, invasive species and climate change. It’s implications are far reaching, and although research in synthetic biology began decades ago, conservation biologists have only recently begun to take notice and appreciate it’s relevance to the conservation of biological diversity. A conference organised by the Wildlife Conservation Society in 2013 discussed the relationship between synthetic biology and conservation, and included speakers from both fields.

Finding Common Ground
It might be surprising to find that, despite a similar background in biological research, the shared knowledge and language of conservationists and synthetic biologists is relatively limited. Further, many synthetic biologists come from an engineering background, with little training in ecology. This can make communication between scientists in these fields more difficult, and may have slowed the pace at which synthetic biology has interfaced with conservation science. The two disciplines also employ different methods and think about nature in different ways. Synthetic biology is largely conducted within large, highly controlled laboratory conditions, whilst ecologists work on complex, interrelated natural systems with a major social and political component. Conservationists, working in a high-stakes field and learning from past mistakes, tend to be quite risk-averse in their practice of conservation, whilst synthetic biologists, working in a new science with much to gain from experimentation, tend to be more in favour of taking large risks. They may also have different outlooks on the future of biodiversity – conservations tend to be more pessimistic about the future, mourning past biodiversity loss, whilst synthetic biologists have an upbeat attitude, envisaging the applications of exciting research. Despite these (extremely generalised) differences, the conference revealed interest and excitement on both sides about the possibility of collaborating, and a mutual appreciation that the major challenges of the Anthropocene are human influences on climate, biodiversity and ecosystems. Finding practical, long-lasting and safe solutions to the plethora of challenges currently facing humanity, is of mutual interest.

Mitigating Risks and Maximising Benefits
The possible applications of synthetic biology to conservation are many. Synthetic biology might enable us to develop more efficient methods of energy production, freeing up habitat to recover. It could mitigate the effects of greenhouse gas emissions by releasing carbon-consuming algae. It could revive extinct species such as mammoths and dodos in a process known as ‘de-extinction’. It could engineer coral that is tolerant to increases in ocean temperature and acidity, conditions which are predicted to worsen under climate change. It could help to control or eradicate invasive species. It could restore degraded land and water for agriculture, sparing the need to destroy more natural habitat. It could even create pesticide- and parasite-resistant bees that can continue to pollinate our crops generations into the future.

However, he potential risks of synthetic biology to conservation are as many as the potential benefits. The effects of synthetic biology on conservation could be direct, (e.g. engineering resistant species), or they could be indirect (e.g. changes in land use). These effects could be negative, for example, if they lead to land use change of primary habitat as has been associated with GMOs and biofuels. They could also be positive, for example if they reduce the impact of human activities, allowing habitat to recover to its natural state. Synthetic biology might lead to unexpected impacts on ecosystem dynamics and risks the unintended escape of novel organisms into open ecosystems. Releasing synthetically engineered organisms into wild environments could alter ecosystems, reduce natural genetic variability or lead to hybridisation events that might display native flora and fauna, and generate new invasive species. Synthetic biology might also distract attention and funds from more traditional conservation efforts, whilst attracting protest from human rights and environmental organisations. Both conservationists and synthetic biologists are conscious of these potential risks, and are committed to careful consideration on a case-by-case basis. Not all synthetic biology is the same; some could be of huge potential benefit to conservation and sustainability whilst carrying minimal risks, and it is these that we should pursue.

Original Article:

() Oryx

Measure Twice, Cut Once:
Quantifying Biases in Sexual Selection Studies

By Claire Asher, on 25 June 2014

Bateman’s principles are conceptually quite simple, but form the basis of our understanding of sexual selection across the animal kingdom. First proposed in 1948, Bateman’s three principles posit that sexual selection is more intense in males than in females for three reasons:

1) males show more variability in the number of mates they have (mating success);
2) males show more variability in the number of offspring they have (reproductive success);
3) the slope of the relationship between mating and reproductive success is steeper in males;

Together, this summarises our basic view of sexual selection in the majority of sexually reproducing species – males that do well, do very well and we expect more intense sexual selection because of it.

Biased Traditions
Traditionally, most studies investigating these relationships have measured mating success by counting the number of females a male produces offspring with. This method is biased though, as it assumes that every mating results in offspring, which is unlikely to be true. Further, it assumes that every fertilisation produces an offspring, which ignores cases where embryos die before birth. Using offspring counts as a way to measure mating success might not be accurate but it is certainly more practical – behavioural observations of actual mating would be very time consuming and nearly impossible for some species. However, until now no study has attempted to quantify the importance of these biases in calculating and testing Bateman’s principles.

Carefully Observed
To address this issue, GEE researchers Dr Julie Collet and Dr Rebecca Dean, in collaboration with researchers at the University of Oxford, University of Queensland, Uppsala University and the University of East Anglia, investigated mating and reproductive success in Red Junglefowl (Gallus gallus). They recorded matings and collected all eggs laid from 13 groupings of 3 males and 4 females (mimicking natural conditions). They began by using classic techniques to estimate Bateman’s gradients – they inferred mating success from the number of females they sired an offspring with. They found twice as much variability in male mating success, and four times as much variance in male reproductive success (the actual number of offspring a male produced) compared with females. Mating success and reproductive success were strongly related – differences between individuals in mating success explained 57% of variance in reproductive success in males, but only 24% in females, and the slope of the relationship was steeper in males.

A Male Red Jungle Fowl

They then repeated their analysis but with a more accurate measure of mating success – the actual number of partners and matings observed. In this study, 30% of pairs that mated did not produce any offspring together and would be ignored by the traditional measure of ‘mating success’. Including these matings reduced the variability in mating success in both males and females. It also reduced the explanatory power of mating success – using this technique they found that variation in mating success actually explained 43% of variation in male reproductive success, and just 5% of variation in female reproductive success. This suggests that traditional methods for measuring Bateman’s principles are likely to be overestimating their importance and the extent of sexual selection on males.

Covarying Factors
Reproductive success is not just a product of how many mates you have. The fecundity of your mate is also a crucial factor, and in species where females mate with multiple males, your share of her offspring is also a key variable. The authors investigated whether these variables tend to be related and whether multivariate analyses that take them all into account better explain the overall reproductive success of a male. Their multivariate model explained the variance in male mating success better than the standard approach and found that mating success, paternity share and mate fecundity together are responsible for the variance in male reproductive success. The authors estimate that by ignoring these other factors, other studies may overestimate the Bateman gradient by as much as 150%!

This study shows the importance of investigating the biases we introduce into our science. These biases may sometimes be inevitable, if excluding them is extremely time consuming or difficult. But we must try to understand the influence of these biases in order to draw informed conclusions from our data. Here, GEE researchers demonstrate how using biased measures of mating success can cause scientists to overestimate the opportunity for sexual selection on males. This effect is likely to be largest for species which have small clutch sizes and in which sperm competition plays a key role. Where possible, studies investigating sexual selection should include accurate measures of mating success, and include other variables such as paternity share and mate fecundity in a multivariate approach in order to best understand Bateman’s principles and the relationship between mating and reproductive success in both sexes.

Original Article:

() Proceedings of the Royal Society B

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This research was made possible by funding from the Natural Environment Research Council (NERC), the Biotechnology and Biological Sciences Research Council (BBSRC) and Marie Curie Action.

Technology for Nature?

By Claire Asher, on 16 June 2014

Many of our greatest technological advances have tended to mark disaster for nature. Cars guzzle fossil fuels and contribute to global warming; industrialised farming practices cause habitat loss and pollution; computers and mobile phones require harmful mining procedures to harvest rare metals. But increasingly, ecologists and conservation biologists are asking whether we can use technology to help nature. On 10th June 2014, UCL’s Center for Biodiversity and Environment Research (CBER) hosted academics from the National Museum of Natural History, Paris, the Zoological Society of London and the Natural History Museum, London for a workshop on “Technology for Nature”. The workshop formed part of a series of public debates and workshops organised in collaboration with the French Embassy, around the theme of the ‘State of Nature’.

Technology for Nature Workshop

The workshop discussed some of the latest technologies available to monitor biodiversity and how these might be harnessed in combination with citizen science to better understand the natural world around us. Citizen science, which engages members of the public in collecting and processing data about nature, is a powerful tool enabling scientists to collect much larger quantities of data on populations of key species. Citizen science projects not only provide biologists with vastly more data than they could ever hope to collect on their own, but it also serves to engage members of the public with the natural world, and raise awareness of key environmental issues.

Following the workshop, UCL also hosted an evening debate in collaboration with colleagues at the French Embassy, London. Professor Romain Julliard from the National Museum of Natural History, Paris, and Professor Kate Jones from UCL’s CBER discussed how new technologies can be used to understand and predict the impact of humans on the natural world, and whether these technologies can be used to inspire and engage the public with the environment around them.

Technology for Nature Debate

Professor Julliard is the Scientific Director of Vigie Nature, a project to monitor trends in various widespread species including butterflies, birds, bees and flowers, using citizen scientists in France. Professor Jones holds the chair of Ecology and Biodiversity at UCL and the Zoological Society of London, and has been involved in a number of projects utilising citizen scientists to monitor populations of bats both in the UK and across Europe. She started the iBats project, which uses volunteers to collect acoustic recordings of bat calls which a computer algorithm can then use to identify the species, and has used citizen science to process data from this project through Bat Detective.

The meeting last week brought together academics from a range of different institutions with a shared interest in monitoring biodiversity to better understand how humans are impacting upon it. We hope this will lead to new projects and collaborations to monitor biodiversity and gain vital data that is needed to assess and ultimately mitigate our impact on the animals and plants we share the planet with.

Nice Flies Don’t Finish Last:
Meiotic Drive and Sexual Selection in Stalk-Eyed Flies

By Claire Asher, on 12 June 2014

While it might seem as though our genes are all working together for our own good, some of them are actually rather selfish. Scientists have known about ‘selfish genetic elements’ for nearly a century, but research to understand their behaviour and effects is ongoing. Recent research in GEE reveals how sexually selected traits are signalling selfish genetic elements (or a lack of them) in the same way they are used to signal male quality and health.

Selfish Genes and the Balance of the Sexes
Selfish genetic elements are variants (alleles) of genes which, rather than acting to the benefit of the individual, act in their own interest to ensure maximum replication of themselves. One type of selfishness that genes can exhibit is called meiotic drive, and is associated with alterations to the sex ratio of offspring. Sex ratios for most species tend towards 1:1, for sound evolutionary reasons, but this isn’t in the best interest of all genes. Genes that lie on one of the sex chromosomes (X or Y in mammals, Z or W in birds) are not equally successful in both sexes – if you happen to lie on the X chromosome, for example, there’ll be two copies of you in each female offspring but only a single copy in male offspring. Meiotic drive occurs when selfish genetic elements skew the sex ratio in order to favour their own replication – usually caused by genes on the X chromosome creating a female-biased sex ratio.

Sex ratios tend to be roughly even in nature because offspring sex ratio is a trait that is under strong stabilising selection. In populations with a biased sex ratio, the underrepresented sex immediately becomes extremely valuable, simply by virtue of its rarity. Imagine a village consisting of 20 women and a single man – that man would undoubtedly have the pick of the ladies, and would likely produce more offspring. If you are the mother of that male, you’ll have done very well for yourself. In a population with a skewed sex ratio, offspring of the underrepresented sex are more valuable, and natural selection to produce them is very powerful. Any mechanism by which females could identify a male that will produce these valuable offspring would be strongly favoured by selection.

Photograph of a male stalk-eyed fly.

A Male Stalk-Eyed Fly (Teleopsis dalmanni)

Meiotic Drive
Stalk-eyed flies are found in Africa and Southeast Asia and show striking sexual dimorphism (physical differences between the sexes). Both males and females have their eyes placed on the end of long stalk-like appendages, but in males these ‘stalks’ can be very long. This is a sexually selected trait – males with longer eyestalks are healthier, carry fewer harmful mutations and are better at attracting females, meaning they tend to have more offspring.

In the laboratory, stalk-eyed flies often produce very female-biased broods; this is thought to be a result of meiotic drive that causes male sperm (carrying a Y chromosome) to degenerate, but female sperm (carrying an X chromosome) to persist. Researchers believe that the length of eyestalks may be linked to meiotic drive, and long-eye stalks may signal to females that a male does not carry the harmful selfish allele. In the laboratory, it has already been shown that males selectively bred for short-eye spans tend to produce female-biased broods, and four loci on the X-chromosome have been identified that are associated with female-bias.

Beyond the Laboratory – Tests in Wild Populations
Expanding on this research, academics in GEE wanted to investigate this phenomenon in the wild. Their work, recently published in Heredity, investigated eyespan and sex ratio biases in 12 populations of wild stalk-eyed flies in Malaysia. Dr Alison Cotton and colleagues at UCL and the University of Debrecen, Hungary, collected nearly 500 wild stalk-eyed flies, measured their eyestalks and other physical characteristics, and collected DNA samples. The researchers used a technique known as microsatellite genotyping to identify regions of the X-chromosome where genes responsible for meiotic drive and for eyestalk length were located. Microsatellites are repeating DNA sequences that tend to vary in their length (the number of repeats) between individuals. Microsatellites can be used as genomic markers for nearby genes of interest – we expect that different microsatellite alleles will be consistently associated with different alleles of interesting genes nearby. Because they vary in physical length, microsatellite alleles can easily be identified by separating DNA sequences out according to size.

The authors found that one microsatellite, ms395, was strongly associated with male eyestalk length. This relationship was not found in females. Longer ms395 alleles tended to be associated with smaller male eyespans.

Photograph of Stalk-eyed flies, (Teleopsis dalmanni)

Stalk-eyed flies, (Teleopsis dalmanni)

Males collected from 5 wild populations were taken back to the lab where they were allowed to mate, so that researchers could investigate the sex ratio of their offspring. Around a quarter of wild-caught males produced biased sex ratios, most often producing more females than males. Males producing sex-biased offspring tended to have smaller eyestalks, when their overall body size was controlled for. They also tended to have longer ms395 alleles.

The authors were able to show that microsatellite ms395 is associated both with sex ratio biases and with male eyestalk length, suggesting that the genes controlling eyestalk length and meiotic drive are located on the X-chromosome near ms395, and that eyespan may be a signal of the genetic quality of the male. For females, a male that carries a selfish genetic element that causes meiotic drive (and a lack of male offspring) is of poor genetic quality, but by preferentially mating with males with longer eyestalks, females can avoid these harmful genes. This may indicate that eyestalks are an example of the good genes hypothesis for sexual selection, which suggests that physical characteristics involved in mate choice are associated with alleles that produce high quality males and are therefore used as an honest signal of the quality of potential mates.

This study is the first to demonstrate meiotic drive in wild populations of the stalk-eyed fly (Teleopsis dalmanni) and adds strong support to previous research suggesting that male eyestalk length is a signal of the presence of meiotic drive. As long as these genes remain in close proximity to each other through evolutionary time, the association should be maintained and eyestalks can be used as a reliable way for females to identify good quality males. Or at least males that are likely to give them a nice even sex ratio in their offspring.

Original Article:

() Heredity

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This research was made possible by funding from the Natural Environment Research Council (NERC), the Engineering and Physical Sciences Research Council (EPSRC) and Marie Curie Action.

Finding a Place to Call Home:
Translocation and the Plight of the Hihi

By Claire Asher, on 16 May 2014

Climate change alters how climate is distributed both geographically and temporally. Over the coming decades, for species sensitive to climatic variables, it may become a case of ‘relocate or die’ – those species that are not able to shift their populations from old, unsuitable habitat into newly emerging suitable habitat, in line with climate change, will likely go extinct. Conservationists can provide a helping hand to species in this position, however – translocation programs aim to establish populations in appropriate habitat when the species is unlikely to reach it on their own. Determining whether translocations are likely to be necessary in the future, what populations to move and where to move them are complex questions to answer, however. Recent work by researchers at the Institute of Zoology (part of the Center for Ecology and Evolution and affiliated with UCL’s GEE department) developed a framework for understanding species’ relationships with climate and identifying potential translocation sites which will provide suitable habitat through future climate change. For one of New Zealand’s endemic birds, the Hihi, translocation to sites further south may be it’s best chance of long-term survival.

Hihi, endangered bird endemic to New Zealand

The climate is changing. Changes in temperature, rainfall and seasonality are occurring globally, and we are already measuring the effects on wildlife. Often, conditions are shifting geographically, and many species will find that their current range no longer overlaps with any suitable habitat (human land-use change isn’t helping!). In these cases, some species will be able to shift their ranges to account for this, but many species will be unable to do change quickly enough to keep up and instead face extinction. Humans can intervene here by moving endangered species to more suitable habitat, but translocation is expensive and it is crucial to select the new location carefully if the population is to have a chance of succeeding. IoZ researchers set out to develop a statistical framework for determining suitable translocation habitat, using one of New Zealands most endearing but endangered endemics, the Hihi (Notiomystis cincta), to test the framework.

The population of Hihis in Tiritiri Mantangi island offers a special opportunity to study the direct effects of climate change without other variables such as food ability confounding the results. This is because they have been provided supplementary food for nearly two decades. Using data on the reproductive success of females in this population, combined with climate data, Dr Aliénor Chauvenet and Dr Nathalie Pettorelli from the Institute of Zoology, along with colleagues at Imperial College London and Massey University in New Zealand, were able to show that Hihi populations are effected by the climate even when food availability is removed from the equation.

Next, using mathematical modelling, the authors tried to predict the future of Hihi populations, using different simulated changes in climate based upon the variables that were found to be most important in influencing current Hihi populations on Tiritiri Mantangi. Changes in temperature, as well as increases in climate variability had a significant influence on the survival of simulated Hihi populations. The final step was to again use mathematical modelling to predict and map suitable Hihi habitat both now, and in the future. Again, this modelling showed that current Hihi populations are most strongly influenced by temperature, a key variable in determining habitat suitability, with rainfall as another important influence.

Looking forward, under models of predicted future climate change, suitable Hihi habitat is expected to move south. The north of New Zealand, which currently offers highly suitable habitat, is predicted to become almost entirely unsuitable over the next few decades. The most successful reintroduced population of Hihis, as well as the largest and last remaining natural Hihi population both stand to lose suitable habitat by 2050. New suitable habitat is expected to emerge in the southern end of the North Island, as well as the northern part of the South Island, where historically conditions have not been suitable for Hihis.

Because Hihis show population declines as temperatures warm even when we control for food availability, even careful management of existing population may prove ineffective under future climate change. Instead, translocation may provide the only solution to guarantee the long-term survival of the Hihi in New Zealand. Although translocations traditionally perform the role of reintroduction – returning a species to part of it’s historical range – future plans for endangered species like the Hihi need to take climate change into consideration. We should opt for ‘assisted colonisation’ – introducing populations to new habitat that is likely to persist (and perhaps even become more suitable) through future climate change. In this way we can attempt to ‘future-proof’ our conservation efforts and hopefully ensure the survival of many species which might otherwise go extinct as the climate changes.

Original Article:

() Journal of Applied Ecology

This research was made possible by funding from AXA Research and Research Councils UK .

Two’s Company, Three’s a crowd:
The Evolution of Two Sexes

By Claire Asher, on 6 May 2014

You’ve probably never given much consideration to why there are men and women. Or, more specifically, why there are two sexes, rather than one, three or 50. But this is a question that has been keeping some scientists awake at night for decades. Recent research in the department of Genetics, Evolution and Environment used mathematical models of evolution to investigate how the evolution of the two sexes was influenced by the inheritance patterns of the energy-producing organelle, mitochondria. The results of this model contradict previous work supporting the idea that inheritance of mitochondria through only one parent might explain the emergence of two sexes. The evolutionary dynamics of mitochondrial inheritance are more complex than previously thought.

Sexual reproduction is a beneficial thing, in evolutionary terms, but this benefit doesn’t depend upon there being different sexes, only on there being two individuals sharing their genes to produce an offspring. This system would also work with no sexes at all (everyone can mate with everyone), or with many sexes. In fact, two is actually the worst number you could have picked – with two sexes any individual is limited to an available pool of mates just 50% of the population. With three sexes, this pool would increase to 66% of the population, with four 75%, and so on. So why have most sexually-reproducing species on settled on two sexes?

In a few previous GEE blog articles (see here and here), I have discussed the phenomenon known as ‘uniparental mitochondrial inheritance’ (UPI), in which mitochondria, organelles found in our cells that are responsible for generating energy, are inherited only through the maternal line – that is, you inherit all of your mitochondria from your mother and none from your father. UPI is found in many living things, although some species do things a bit differently and there are many different ways to achieve the same result. Work by GEE researcher Professor John Allen has previously shown that the mitochondria within egg cells in jellyfish, fruit flies and fish are largely inactive; this inactivity allows for a perfect ‘mitochondrial template’ to be passed on to the offspring and prevent the accumulation of mutations through the generations. Essentially, this is why aging isn’t heritable. It wouldn’t work to inactivate sperm mitochondria because they need so much energy for all that swimming, so if we did inherit mitochondria from our fathers they would probably be mutated. UPI is also thought to help evolution remove harmful mutations from the population and reduce conflict and promote coadpatation between the mitochondrial symbiont and its host cell.

Fertilisation So, UPI makes a lot of sense, evolutionarily, and some scientists think it might also explain why we have two sexes, as opposed to any other mating system. It’s important to be clear, when we talk about having two sexes we’re saying nothing about the external differences between the sexes (sexual dimorphism) observed in many multicellular organisms. We’re talking about the existence of two ‘mating types’, such that individuals cannot mate with members of the same type. Recent research by another group of GEE academics including Professor Andrew Pomiankowski, Dr Nick Lane and Professor Robert Seymour, investigated the evolution of UPI and in particular it’s relationship with the evolution of a two-sex mating system. We might expect a strong link between UPI and the existence of two sexes, since uniparental inheritance immediately generates differences between the two mating partners, and ensures that reproduction is not possible unless one member of each ‘type’ is present. Although UPI is often thought to have been a key driver in the evolution of mating types, there have been few investigations of what conditions are needed for the fitness benefits of UPI to actively drive the emergence of two mating types. So the authors developed a new mathematical model of mitochondrial inheritance and the evolution of UPI in a population where biparental inheritance (BPI) is the norm. They incorporated mitochondrial mutation (which might sometimes be selfish) and selection into the model, and included different mating types.

The model agreed with a great deal of previous work that indicates that UPI tends to increase fitness. It does so slowly, with selection acting cumulatively across many generations to remove less fit mitochondrial variants and increase fitness for UPI individuals. In a population of individuals where mitochondria is inherited biparentally, a new mutation causing UPI exists in a single individual. Slowly UPI improves the fitness of cells by reducing the number of mutated mitochondria they carry, and the UPI mutation might start to spread in the population. The problem is, as it spreads the benefits of UPI are inevitably leaked into the rest of the BPI population – UPI individuals mate with BPI individuals producing some BPI offspring who carry the fitter mitochondria from their UPI parent. This leaking of benefits means that the fitness benefits of UPI are frequency-dependent; the more common UPI becomes in a population, the less each UPI individual benefits from the reproductive strategy. This makes it hard for UPI to fully take over a population – their model tended instead to produce mixed populations with some UPI and some BPI individuals interbreeding.

Mitochondrial Evolution - Leaking of beneficial mutatations

Leaking of fit mitochondrial (blue) into BPI cells (a)

If the researchers included mating types in the model at the start of it’s evolutionary run, then UPI could become associated with specific mating types and in this situation, so long as mutation rates were high or each cell carried many mitochondria, UPI could spread to fixation in the population. But UPI itself was not able to alter the number or existence of mating types. The authors suggest that this may explain the continuum of UPI levels we observe in nature. For any given species, the occurrence of UPI will depend upon the evolutionary starting point, energetic demands, mutation rates and the selfish (or unselfish) nature of mutations.

Although most people never even consider why we have two sexes, male and female, the evolution of a two mating-type system is seemingly paradoxical and many theories and hypotheses have been proposed to explain it. One such explanation is that uniparental inheritance, which is critical for stabilising the mitochondria-cell symbiosis and preventing the accumulation of harmful mutations, may have driven the evolution of two sexes. However, mathematical modelling by scientists in GEE suggests this is not the case, and UPI more likely evolved after the two mating-type system emerged. In their model, although UPI initially spreads through populations, it’s fitness benefits are frequency-dependent, meaning it only rarely takes over an entire population. Populations in which all members inherit mitochondrial uniparentally are only possible when a mutation causing UPI becomes tighly linked to genes that determine mating type. The initial emergence of two mating types still requires an explanation independent from mitochondrial inheritance patterns.

Original Article:

() Proceedings of the Royal Society B: Biological Sciences

This research was made possible by funding from the Natural Environment Research Council (NERC), the Engineering and Physical Sciences Research Council (EPSRC) and the Leverhulm Trust.

Size Matters for Single-Celled Dating

By Claire Asher, on 24 April 2014

Size matters. Particularly when it comes to finding a mate. Across the animal kingdom, body size plays a crucial role in determining how successful individuals are at surviving and reproducing. Large males are often better at fighting off competing males, while large females generally produce more offspring. However, being big isn’t always best. In some environments, small individuals may have a better chance of survival. Choosing a partner the right size may help to ensure a good start in life for your offspring. Recent research in GEE suggests that even simple organisms such as yeast are able to choose the best size mate to ensure their offspring survive and reproduce.

In single-celled organisms, just like their multicellular counterparts, the ideal body size depends on surrounding conditions in the environment. In nutrient rich environments, larger cells do better, whilst in nutrient poor environments, smaller cells prevail. In many multi-cellular animals, females have been shown to select mates based on body size to ensure their offspring the best possible chances of survival. Are unicellular organisms also able to make these kinds of selections? A recent paper by GEE Researchers Dr Carl Smith, Prof. Andrew Pomiankowski and Dr Duncan Greig investigated this phenomenon in yeast.

The Curious Life-Cycle of Yeast
Yeast Life CycleYeast, being single-celled, are able to reproduce asexually – budding off a new copy of themselves. However, when times are hard, yeast can also enter a sexual phase of reproduction. When nutrient levels are low, a yeast cell will divide into sexual spores. These spores are haploid, carrying only one copy of each chromosome (rather than the usual two), making them analogous to sperm and egg cells. Spores will remain dormant until they sense an improvement in the environment, at which point they spring into action and try to find a mate. Spores come in two mating types and must always mate with another spore of the opposite type – a system based on mating pheromones released by spores to attract the opposite sex.

Body size is important in both the asexual and sexual phases of yeast reproduction. Yeast cells must reach a certain critical size before they can bud asexual copies of themselves, although this critical size is smaller in poorer environments. Likewise, during sexual reproduction, the cell size of the offspring is determined by the cell size of the two spores that fuse together. GEE researchers first investigated how body size influenced reproductive success in both phases of yeast reproduction. Cell size was an important influence on how quickly spores awoke from dormancy to begin searching for a sexual partner; large spores became active 38% more quickly in a nutrient-rich environment, whereas smaller spores were 63% faster when conditions were poor. Asexual reproduction was similarly affected, with large spores budding off asexual copies sooner in good environments and small spores budding sooner in bad environments. Size matters when it comes to yeast reproduction, but bigger isn’t always better.

Are Sexual Yeast Cells Selecting the Right Sized Partner for their Environment?
To answer this question, GEE Researchers created a mating-choice experiment, in which a haploid yeast cell of one mating type is presented with two cells of the opposite type, one small and one large. To mate, two of these haploid cells will fuse to form a single, diploid cell which can resume asexual reproduction. These experiments were performed with different sizes of focal yeast cell, and in different environments, to see whether the focal cells were able to select the best partner – one that would give the final fused cell the best chance of survival. This is a critical choice – mating in these single-celled organisms is an irreversible decision, the size of the ‘offspring’ cell is directly determined by the size of the two haploid cells that fuse, and cell size will influence how quickly the ‘offspring’ can reproduce.

Yeast

The results of the experiment indicate that yeast cells are indeed able to make careful calculations about their choice of partner and select the one closest to the optimum size for the environment. In nutrient rich environments, yeast cells tended to select larger mates, whereas in poorer environments they tended to select smaller mates. This was true regardless of their own body size, indicating that cells were not simply mating with similar sized partners, but making a choice based upon the mates available and the environment around them. The authors suggest that such seemingly complex behaviour could be generated quite simply – if cells that are better adapted to their environment begin emitting these pheromones earlier, or produce them in larger amounts, they would be more easily detected by a prospective mate.

Size is a crucial determinant of success, survival and reproduction for most living organisms. However, the optimum size often depends upon environmental conditions, as GEE researchers have shown to be the case for yeast. For most organisms, selecting a mate is an important decision that will influence the success of their offspring. For yeast even more so, this decision is critical, as it is irreversible. A new paper from GEE researchers shows that yeast cells also consider the quality of their partner when mating. During sexual reproduction, yeast cells are able to select a mate of the optimum size for the environment, thereby improving the chances that the resulting offspring cell will reproduce.

Original Article:

() Ecology and Evolution

This research was made possible by funding from the Natural Environment Research Council (NERC), the Biotechnology and Biological Sciences Research Council (BBSRC), the Engineering and Physical Sciences Research Council (EPSRC), The Royal Society and the Max Planck Society.

Dating Mammalian Evolution

By Claire Asher, on 28 March 2014

When the age of the dinosaurs ended around 65 million years ago, mammals stepped in to fill the gap, and the age of the placentals began. However, whether early placental mammals were already present on Earth before the demise of the dinosaurs has been the subject of a long standing debate. Recent research in GEE used genomic data, in combination with fossil evidence, to show that the earliest placental mammals were indeed scurrying between the feet of dinosaurs.

Shrew-like mammals scurry between the feet of dinosaursThe huge diversity of placental mammals on Earth today first appeared shortly after the mass extinction event that killed the dinosaurs. It is thought that the loss of the dinosaurs, along with much of life on Earth, freed up niches which placental mammals to evolved to fill. But were early placental mammals present, waiting in the wings, during the age of the dinosaurs, or did they appear rapidly after their demise? One recent study suggested that, based on fossil evidence, the placental mammals must have appeared after the cretaceous-tertiary boundary (KT) when dinosaurs and most life on Earth was wiped out. However, a recent paper by GEE’s Mario dos Reis and Ziheng Yang, in collaboration with Philip Donoghue from the University of Bristol, highlights flaws in the methods used in this study, and utilitsed a more thorough approach to show that early placental mammals likely predated the KT boundary.

Using genetic sequence data from over 14,000 genes, combined with fossil evidence, GEE researchers applied 3 alternative statistical methods to estimate the age of the earliest placental mammal; ancestor to all modern placental mammals. Although different statistical methods yielded slightly different estimates, and differed in their accuracy, they all agreed that placental mammals must have already been around before the dinosaurs went extinct. The adaptive radiation of mammals that occurred after the extinction of the dinosaurs was dramatic, but it was initiated by a few shrew-like species which had already evolved. This study highlights the importance of using statistical methods to estimate the true age of ancestral species; the age of the oldest fossils is not the same as the age of the ancestral species that gave rise to them, and statistical techniques must be employed to estimate this. Using both molecular and fossil evidence to inform estimates also provides more robust evidence for the true age of the first placental mammals, and the theory that the earliest ancestors of placentals predated the disappearance of the dinosaurs.

Original Article:

This research was made possible by funding from the Biotechnology and Biological Sciences Research Council (BBSRC).

The Delicate Balance of Effect and Response

By Claire Asher, on 18 February 2014

We may not always be aware of it, but many wild plants, animals, fungi and even bacteria, provide crucial services to us which keep the ecosystems of Earth functioning. Environmental changes caused by human activities are now threatening many species, and those that cannot withstand these changes may be lost forever, potentially taking the services they provide away. New research from GEE and collaborators worldwide aims to improve our understanding of how the traits and evolutionary histories of species influence their ability to provide essential ecosystem services, and to persist in the face of ongoing environmental change.

The diverse array of species we share planet Earth with, and the complex ecosystems they form, are crucial to our continued survival and well being. Species and ecosystems provide a huge number of ‘ecosystem services’ – functions such as nutrient cycling, waste decomposition, pollination and food, to name just a few, which humans rely on. However, many species are now under threat from human activities like deforestation, hunting and pollution. Scientists are working to understand how species and ecosystems will respond to our continued activities in the future, and particularly how this may effect the vital ecosystem services upon which we rely. Recent research by GEE’s Prof. Georgina Mace, in collaboration with researchers from Cordoba National Univerity, Imperial College London, VU University, Yale University and CSIC, has attempted to develop a new framework for risk assessing the effect of human activities on ecosystem services.

The framework considers two key aspects of species: their effect on the generation of a specific ecosystem service (e.g. seed dispersal), and their response to specific environmental pressures (e.g. drought). Both the effect of a species and the response of a species are underpinned by its traits, and each is influenced not by a single trait but a combination of traits. The response of a species will determine it’s ability to survive and flourish through future environmental changes and to continue to provide it’s ecosystem effects. However, only a species’ response is the subject of natural selection, via changes to the underlying traits; the effect of a species is merely a biproduct of traits selected for their influence on survival. In this way, the aspects of a species’ biology upon which we rely are only indirectly influenced by natural selection, and will only be maintained if the traits that generate them are beneficial through the environmental changes we cause. The framework developed by GEE researchers and collaborators considers how the response of species to envinmental stressors interacts with the effect of that species on key ecosystem services, and whether species with a large effect are more or less vulnerable to environmental change.

A third key factor influencing the sensitivity of ecosystem system services is the evolutionary relationships between species providing them. Closely related species often share similar traits, which may or may not result in them having similar effects and responses. If this is the case, then ecosystems in which a particular service is provided by a group of closely related species may be more vulnerable to environmental change, since those species may well share similar responses, and be sensitive to similar environmental pressures. Although many species’ traits are known to be similar amongst related species, because effects and responses are each the result of a combination of traits, it is not known whether this relationship is also common for these variables.

The new framework developed by GEE’s Professor Georgina Mace and collaborators attempts to address this by incorporating evolutionary relationships (phylogeny) into their response-effect model, and applying this model to 5 case studies. The case studies cover 5 species assemblages including a total of 480 species in Europe, Central America and Africa, for which response and effects could be estimated based on past studies of species’ traits and vulnerabilities. The case studies tended to show a strong relationship between phylogeny and both species’ effects on ecosystem services and their responses to environmental stressors. This indeed suggests that ecosystem services that rely upon closely related groups of species may be most at risk from environmental change. Cases where effects and responses are negatively correlated, so that the most influential species in terms of a given ecosystem service are also the most vulnerable to environmental stress, are most vulnerable to loss of that ecosystem service through human activities. Whether this type of relationship is common in nature remains to be investigated by future studies, and this framework provides a powerful basis with which to do so.

Our relentless demands on the natural world are inevitably leading to new pressures and stresses on natural populations, and it is of great concern that these pressures may negatively impact on the vital ecosystem services that we rely upon, often without even realising it. Ecosystem services provide us with food and fresh water, decompose our waste, recycle nutrients and remove harmful toxins. Without them our continued survival and well being would be seriously compromised. Scientists are still working to understand how species’ traits influence their ability to provide ecosystem services and their resilience to ongoing environmmental change. A new framework developed in collaboration between universities in the UK, Spain, Argentina, the USA and the Netherlands is beginning to shed light on the interaction between species’ traits, their effect on ecosystem services and their response to environmental change, and how these factors are influenced by evolutionary relationships between species. This framework offers a powerful new view of how the traits of species within an ecosystem translate into the ecosystem services upon which we are so reliant, and future research building upon this framework promises to improve our understanding of ecosystem services and environmental change.

Original Article:

() Ecology and Evolution



This research was made possible by funding from the Natural Environment Research Council (NERC), the Leverhulme Trust, and the US National Science Foundation