Tag Archives: Research Blogging

“Chemical camouflage” lets leafhoppers hide from their own bodyguards

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ResearchBlogging.orgMany insects in the order Hemiptera – the “true” bugs – have evolved a way to hire their own protection by excreting sugary “honeydew.” Honeydew attracts ants, who tend honeydew-producing bugs like livestock, protecting them from predators and even disease. Honeydew is cheap to make because honeydew producers typically make a living sucking the sap of their host plants; they’re trading sugar and water, which they have in abundance, for safety.


Camponotus crassus ants protect Guayaquila xiphias leafhoppers, apparently mistaking them for part of their host plant. Detail of Silveira et al., figure 1.

But there’s a catch. Ants make good bodyguards because they are carnivorous – and they’re perfectly willing to start eating their flock. A natural history note in the latest issue of The American Naturalist suggests that one group of ant-protected bugs deals with this problem by cloaking themselves in chemicals that make the ants think they’re part of the host plant [$a].

The study’s authors determined that organic compounds on the cuticle of the honeydew-producing leafhopper Guayaquila xiphias, which is often tended by the ant Camponotus crassus, were similar to compounds on the surface of the leafhoppers’ preferred host plant. They presented ants with freeze-dried leafhoppers whose cuticles were washed clean with solvent, and found that the ants were much more likely to attack washed leafhoppers than unwashed ones; the ants were also more likely to attack leafhoppers they found on plants other than the preferred host. Finally, the authors replicated the earlier experiments using moth larvae coated with leafhopper cuticle compounds, and found that the “chemical camouflage” conferred the same protection on a different insect species.

This neat result shows how hazardous honeydew-producers’ relationship with their ant bodyguards can be – they have to hide from the ants even as they offer them an inducement to stick around!

Reference

Silveira, H., Oliveira, P., & Trigo, J. (2010). Attracting predators without falling prey: Chemical camouflage protects honeydew‐producing treehoppers from ant predation The American Naturalist, 175 (2), 261-8 DOI: 10.1086/649580

Evolving from pathogen to symbiont

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This post was chosen as an Editor's Selection for ResearchBlogging.orgRecently the open-access PLoS Biology published a really cool study in experimental evolution, in which a disease-causing bacterium was converted to something very like an important plant symbiont. The details of the process are particularly interesting, because the authors actually used natural selection to identify the evolutionary change that makes a pathogen into a mutualist.

Life as we know it needs nitrogen – it’s a key element in amino acids, which mean proteins, which mean structural and metabolic molecules in every living cell. Conveniently for life as we know it, Earth’s atmosphere is 78% nitrogen by weight. Inconveniently, that nitrogen is mostly in a biologically inactive form. Converting that inactive form to biologically useful ammonia is therefore extremely important. This process is nitrogen fixation, and it is best known as the reason for one of the most widespread mutualistic interactions, between bacteria capable of fixing nitrogen and select plant species that can host them.


Clover roots, with nodules visible (click through to the original for a nice, close view. Photo by oceandesetoile.

In this interaction, nitrogen-fixing bacteria infect the roots of a host plant. In response to the infection, the host roots form specialized structures called nodules, which provide the bacteria with sugars produced by the plant. The bacteria produce excess ammonia, which the plant takes up and puts to its own uses. The biggest group of host plants are probably the legumes, which include the clover pictured to the right, as well as beans – this nitrogen fixation relationship is the reason that beans are the best source of vegetarian protein, and why crop rotation schemes include beans or alfalfa to replenish nitrogen in the soil.

For the nitrogen-fixation mutualism to work, free-living bacteria must successfully infect newly forming roots in a host plant, and then induce them to form nodules. The chemical interactions between bacteria and host plant necessary for establishing the mutualism are pretty well understood, and in fact genes for many of the bacterial traits, including nitrogen-fixation and nodule-formation proteins thought to be necessary to make it work are conveniently packaged on a plasmid, a self-contained ring of DNA separate from the rest of the bacterial genome, which is easily transferred to other bacteria.

This is exactly what the new study’s authors did. They transplanted the symbiosis plasmid from the nitrogen-fixing bacteria Cupriavidus taiwanensis into Ralstonia solanacearum, a similar, but disease-causing, bacterium. With the plasmid, Ralstonia fixed nitrogen and produced the protein necessary to induce nodule formation – but host plant roots infected with the engineered Ralstonia didn’t form nodules. Clearly there was more to setting up the mutualism than the genes encoded on the plasmid.


Wild-type colonies of Ralstonia (tagged with fluorescent green) are unable to enter root hairs (A), but colonies with inactivated hrcV genes are able to enter and form “infection threads,” like symbiotic bacteria (B). Detail of Marchetti et al. (2010), figure 2.

This is where the authors turned to natural selection to do the work for them. They generated a genetically variable line of plasmid-carrying Ralstonia, and used this population to infect host plant roots. If any of the bacteria in the variable population bore a mutation (or mutations) necessary for establishing mutualism, they would be able to form nodules in the host roots where others couldn’t. And that is what happened: three strains out of the variable population successfully formed nodules. The authors then sequenced the entire genomes of these strains to find regions of DNA that differed from the ancestral, non-nodule-forming strain.

This procedure identified one particular region of the genome associated with virulence – the disease-causing ability to infect and damage a host – that was inactivated in the nodule-forming mutant strains. As seen in the figure I’ve excerpted above, plasmid-bearing Ralstonia with this mutation were able to form infection threads, an intermediate step to nodule-formation, where plasmid-bearing Ralstonia without the mutation could not. Clever use of experimental evolution helped to identify a critical step in the evolution from pathogenic bacterium to nitrogen-fixing mutualist.

References

Amadou, C., Pascal, G., Mangenot, S., Glew, M., Bontemps, C., Capela, D., Carrere, S., Cruveiller, S., Dossat, C., Lajus, A., Marchetti, M., Poinsot, V., Rouy, Z., Servin, B., Saad, M., Schenowitz, C., Barbe, V., Batut, J., Medigue, C., & Masson-Boivin, C. (2008). Genome sequence of the  beta-rhizobium Cupriavidus taiwanensis and comparative genomics of rhizobia. Genome Research, 18 (9), 1472-83 DOI: 10.1101/gr.076448.108

Gitig, D. (2010). Evolving towards mutualism. PLoS Biology, 8 (1) DOI: 10.1371/journal.pbio.1000279

Marchetti, M., Capela, D., Glew, M., Cruveiller, S., Chane-Woon-Ming, B., Gris, C., Timmers, T., Poinsot, V., Gilbert, L., Heeb, P., Médigue, C., Batut, J., & Masson-Boivin, C. (2010). Experimental evolution of a plant pathogen into a legume symbiont. PLoS Biology, 8 (1) DOI: 10.1371/journal.pbio.1000280

Why make your own food when it doesn’t pay?

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ResearchBlogging.orgWe humans like to think we’re pretty complex – what with having invented the wheel, wars, New York, and so on – so we tend to forget that evolution doesn’t care about complexity. All that matters to natural selection is who makes the most babies, and sometimes complex adaptations can get in the way of that criterion. A study recently published on the always open-access PLoS ONE provides a good example of this principle in action – given the right selective pressures, photosynthetic organisms will give up on the whole photosynthesis thing.



Tiny Indianpipe (Monotropa) and giant Rafflesia, two plants that gave up photosynthesis. Photos by Bemep and Tamara van Molken.

Photosynthesis is clearly a complex adaptation, requiring specialized cellular structures and biochemical processes that can use light to power the synthesis of sugars. Complex enough for a whole additional organism, in fact, since the chloroplast, the cellular structure in which most eukaryotes conduct photosynthesis, probably originated as a symbiont that never left its host cell [$a]. (In some organisms, this process of becoming photosynthetic is still underway.) There are clear advantages to the ability to make your own food conferred by photosynthesis. Yet there are numerous examples of non-photosynthetic organisms with photosynthetic ancestors. For instance, plants as varied as the big, exotic Rafflesia or Monotropa, whose small white flowers are easy to spot in North American woods, have inactive chloroplasts and parasitize other plants. These cases are good reason to think that there may be selective conditions in which the cost of maintaining the mechanisms of photosynthesis outweighs the benefit of independent food production.

The new paper describes just such a set of selective conditions. The authors build a mathematical model of competition between microorganisms, such as flagellates, that can either be mixotrophs, able to conduct photosynthesis or capture prey to feed themselves, or heterotrophs, only able to sustain themselves by eating other critters. The model’s result hinges on two key facts of life for single-celled predators: (1) it turns out that the size of a flagellate cell determines what size of prey it is best able to capture [$a]; and (2) chloroplasts take up space in a cell, limiting the evolution of cell size.

The relative advantage of retaining photosynthesis, then, is directly related to the size range of available prey. Mixotrophs, whose cells are big enough to accommodate chloroplasts, are most efficient predators of larger prey; with no chloroplasts, heterotrophs can be small enough to take advantage of smaller prey. The question of which form wins out, then, relies on the distribution of available prey sizes and the light environment. If there’s lots of light for photosynthesis, mixotrophs can out-compete heterotrophs even if they don’t hunt very efficiently; but if there’s not much light and mostly small prey, the more efficient heterotrophs win.

The fact is, it’s rare for any given adaptation to be useful under all possible conditions. Biological structures or metabolic processes that become disused are no longer under selection for efficient performance of their original function – they are free to accumulate mutations that may make them degenerate into uselessness, or to be co-opted for entirely new functions. But if an adaptation is actually costly to maintain, then natural selection may eradicate it altogether.

References

de Castro, F., Gaedke, U., & Boenigk, J. (2009). Reverse evolution: Driving forces behind the loss of acquired photosynthetic traits. PLoS ONE, 4 (12) DOI: 10.1371/journal.pone.0008465

Hansen, B., P. K. Bjornsen, & P. J. Hansen (1994). The size ratio between planktonic predators and their prey.
Limnology and Oceanography, 39, 395-403

McFadden, G. (2001). Chloroplast origin and integration Plant Physiology, 125 (1), 50-3 DOI: 10.1104/pp.125.1.50

Masquerading caterpillars hide in plain sight

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ResearchBlogging.orgInsects that have evolved elaborate mimicry of inanimate objects – leaves, twigs, even bird droppings – to hide from predators are a staple of nature documentaries. But do these masquerades work because they help insects blend into the background, or because predators actually see the insects and then dismiss them as inedible leaves, twigs, or bird droppings? It’s a tricky question to answer, but a brief paper in this week’s Science presents an experiment that tries to do just that [$a].

The paper’s authors reasoned that if mimicry-based camouflage works through disguise rather than invisibility, a predator’s experience might determine their response to mimic camouflage. They trained three experimental groups of young domestic chicks by introducing them into trial arenas containing either natural hawthorn branches, empty arenas, or hawthorn branches wrapped in purple thread. The wrapped branches were used to test whether the chicks would be more or less likely to attack something twig-like but differently colored (though this is only clear from the supplementary online material).


Larva of the brimstone moth Opisthograptis luteolata, looking distinctly twig-like. Photo by Michael E. Talbot.

The authors then presented chicks from each “training” group with either one of two species of hawthorn-twig-mimicking moth larvae (the brimstone moth, or the early thorn moth), or a hawthorn twig about the size of a caterpillar. Chicks that had previously encountered natural twigs waited longer to attack the caterpillars than chicks that hadn’t previously seen twigs, or that saw the colored hawthorn branches. So, apparently, the chicks were reasoning (inasmuch as chicks reason) that the twig-like object in front of them was the same as the inedible twigs they had tried before.

This is an elegant experimental test of the effect of mimicry as mimicry – what the authors propose to call camouflage by “masquerade.” However, it doesn’t actually show that what the authors term camouflage by crypsis – blending into the background – isn’t also contributing to the benefits that these caterpillars receive from their unique shape and coloration. There’s no reason to think that twig-shaped caterpillars can’t benefit in both ways, by being less visible in the first place, and then easily mistaken for a twig if they are seen.

In conclusion, here’s some video footage of another natural mimic, the leaf insect.

Reference

Skelhorn, J., Rowland, H., Speed, M., & Ruxton, G. (2010). Masquerade: Camouflage without crypsis Science, 327 (5961), 51 DOI: 10.1126/science.1181931

Escaping the “poverty trap” of infectious disease

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ResearchBlogging.orgEven in the twenty-first century, infectious diseases such as malaria, dengue fever, cholera, and AIDS remain widespread in much of the developing world, at tremendous cost to human life and economic productivity. Poorer nations lack the resources for more effective public health measures; but widespread infectious disease may slow or prevent the economic development that can provide those resources. A new paper in Proceedings of the Royal Society tries to sort out this chicken-and-egg problem, and finds that economic development is the fastest route out of the “poverty trap” [$a].

The paper’s authors, Bonds et al. start with a classic model of infectious disease, in which susceptible (healthy) members of a population have a chance of becoming infected whenever they encounter an infected person, and infected people have a chance to recover to susceptible condition if they survive the effects of the disease. The first probability is the rate of transmission from person to person; the second is the rate of recovery from disease caused by the infection.


A woman receives tetanus vaccine in the Central African Republic. Photo by hdptcar.

Bonds et al. insert economics into this basic model by reasoning that the rate of recovery is a function of per-capita income – well-fed people are better able to fight off infection – and that income is a function of the proportion of the population that remains uninfected at any given time. This yields a mathematical version of the poverty trap I outlined above: high-income populations are easily able to fight off infection and remain near 100 percent susceptible, but highly infected societies are unable to increase their per-capita income to reduce their rate of infection. However, there is an internal equilibrium point – a level of income and infection from which a population could easily move in either direction, towards high income and low infection or high infection and low income.

The question then becomes how best to push a developing nation’s population toward that threshold condition – or how best to bring the threshold closer. Bonds et al. compare two options: reducing the rate of disease transmission, and boosting individual economic productivity. The former captures the effect of boosting public health – vaccination, better sewage treatment, food aid. The latter captures the effect of improving infrastructure or financial institutions – making the economy more developed. They found that the threshold condition is more sensitive to economic productivity. Even at low transmission rates, a society can be caught in the poverty trap if its productivity is low enough, but at high enough productivity levels, societies can avoid the trap created by even highly transmissible diseases.

This suggests that, although medical aid can help the acute problem of infectious disease, it’s investment in economic development that can ultimately solve it.

Reference

Bonds, M., Keenan, D., Rohani, P., & Sachs, J. (2009). Poverty trap formed by the ecology of infectious diseases Proc. R. Soc. B DOI: 10.1098/rspb.2009.1778

Why aren’t there more sickle-cell anemics in the Mediterranean?

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This post was chosen as an Editor's Selection for ResearchBlogging.orgThe story of sickle-cell anemia and its malaria-protective effects is a textbook case how environmental context determines the fitness of a given genetic profile. However, the evolution of human blood disorders in response to selection from malaria parasites might be more complicated than that textbook story.



Malaria-causing parasites (dark-stained) among human red blood cells (top), and “sickled” red blood cells (bottom). Photos via WikiMedia Commons.

Malaria is caused by mosquito-spread parasites that attack their hosts’ oxygen-bearing red blood cells. A particular mutation in the gene that codes for part of the hemoglobin molecule – the molecule that actually stores oxygen inside red blood cells – leads to deformed, sickle-shaped, blood cells. People who carry two copies of the sickle cell gene develop sickle-cell disease, in which the sickle-shaped cells reduce oxygen transport efficiency and interfere with blood circulation. People with only one copy of the sickle-cell gene are healthy, and better able to resist malaria infection than those with no copies. The textbook story is that, in regions where malaria is common, such as sub-Saharan Africa, the advantage of malaria resistance is enough to offset the fitness risk of carrying the sickle-cell gene – that one-fourth of children born to parents who each have one copy of the gene will themselves have two copies and develop sickle-cell disease.

However, there are regions like the Mediterranean where malaria has historically been prevalent, but in which the human population hasn’t evolved the higher frequency of sickle-cell genes that you’d expect from the scenario outlined above. A new paper in PNAS demonstrates that this may be because of interactions between the sickle-cell gene and another genetic blood disorder, thalassemia [$a].

Thalassemia is a class of genetic disorders affecting the protein subunits that comprise hemoglobin. Each hemoglobin molecule is formed by binding together two “alpha”-type subunits, and two “beta”-type subunits. If there is a shortage of correctly-formed subunits of either type, then hemoglobin formation is impaired, resulting in anemia or (if the mutation stops subunit production altogether) death. However, like sickle-cell genes, thalassemic mutations can confer resistance to malaria; and if alpha-thalassemia is paired with beta-thalassemia, the reduced production of both subunits can balance out.

As it happens, in combination with alpha-thalassemia, the sickle-cell gene’s malaria protection is neutralized. Using population genetic models, the new study’s authors show that this effect may have actively prevented the sickle-cell gene from establishing in the Mediterranean, where alpha- and beta-thalassemias are more common than in Africa. In the Mediterranean, the presence of beta-thalassemia genes reduces the fitness cost of (mild) alpha-thalassemia genes; and in the presence of alpha-thalassemia genes, the sickle-cell gene confers no protection to people with one copy but still induces sickle-cell disease in people with two copies.

These interactions between genes are called epistasis, and they can have dramatic impacts on evolution. Although I haven’t seen many cases as well-characterized as this one, epistasis is probably widespread in the complex systems of genomes, where thousands of regulatory and protein-coding genes interact to build living things.

References

Penman, B., Pybus, O., Weatherall, D., & Gupta, S. (2009). Epistatic interactions between genetic disorders of hemoglobin can explain why the sickle-cell gene is uncommon in the Mediterranean Proc. Nat. Acad. Sci. USA, 106 (50), 21242-6 DOI: 10.1073/pnas.0910840106

Cuckholding crows don’t necessarily have healthier chicks

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ResearchBlogging.orgBirds are bad at monogamy. There are a number of good evolutionary reasons to cheat on your mate, and it’s not clear which one is the most likely explanation. A new study of American crows, however, suggests that, for females, cheating isn’t necessarily the best choice [$-a].

Avian infidelity isn’t obvious, because many birds are socially monogamous, forming couples for one or more breeding seasons to raise chicks. However, DNA-based paternity testing has overturned this intuition — a 2002 review of such studies [PDF] estimated that “cheating” occurs in 90% of bird species, and an average of 11% of chicks are “illegitimate.”

The biological term for this non-monogamy is “extrapair copulation,” often abbreviated to EPC. Evolutionary reasons for EPC behavior break down by which parent benefits from the cuckoldry: Females benefit if EPC means their chicks will be less inbred, which can make them less prone to disease or recessive genetic disorders. Males benefit if EPC means they will have more chicks than they would otherwise. Perhaps more importantly, EPC might impose real costs on females, if it leads mated males to invest less in caring for the chicks in their nests because they can’t be sure the chicks are theirs [PDF].


Crows in flight. Photo by wolfpix.

In the new study, Townsend et al. evaluated the costs and benefits of EPC for female American crows, which have a social structure that adds a twist to the cost-benefit analysis. Mated pairs of crows live in larger family groups, which include “auxiliary,” unmated males who may help feed and protect chicks — perhaps especially if those chicks are the result of their own EPC. Females also engaged in EPC with males from outside the family group, who should be less closely related than within-group males, and whose chicks would be more genetically healthy than those sired by any within-group male, mated or not.

Townsend et al. observed several such family groups over four years, using DNA fingerprinting methods to identify the parents of chicks as they were born, and tracking the chicks’ health and survival as well as how frequently mated crows and auxiliary males tended them. Contrary to what might have been expected, chicks produced by EPC were more, not less, inbred; they didn’t grow faster or have a higher probability of survival than chicks produced by mated parents. On the other hand, cuckholded males tended chicks sired by others as often as they did their own.

The most telling result is that broods containing chicks produced by EPC were more frequently tended by auxiliary males — but only when the EPC was with a within-group male. This suggests that EPC mainly benefits male crows, not females. From a mated female’s perspective, EPC produces chicks that are less genetically fit, and no more or less likely to survive, than chicks sired by her mate. On the other hand, an unmated male can only have offspring through EPC, and if he does, it makes sense for him to give them extra assistance. Males from outside the family group don’t stick around to offer that help, but auxiliary males from within the group can, and do.

References

Arnqvist, G., & Kirkpatrick, M. (2005). The evolution of infidelity in socially monogamous passerines: The strength of direct and indirect selection on extrapair copulation behavior in females. The American Naturalist, 165 (s5) DOI: 10.1086/429350

Griffith, S.C., Owens, I.P.F., & Thuman, K.A. (2002). Extrapair paternity in birds: A review of interspecific variation and adaptive function. Molecular Ecology (11), 2195-212 : 10.1046/j.1365-294X.2002.01613.x

Townsend, A., Clark, A., & McGowan, K. (2010). Direct benefits and genetic costs of extrapair paternity for female American Crows (Corvus brachyrhynchos). The American Naturalist, 175 (1) DOI: 10.1086/648553

Picky eating, not genetics, splits leaf beetles

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This post was chosen as an Editor's Selection for ResearchBlogging.orgMany different factors can conspire to create reproductive isolation between populations and, ultimately, separate species. Disentangling them is often tricky, but a study recently published in PNAS takes a crack, and demonstrates that two populations of leaf beetles are divided by food preferences, not genetics [$-a]




Neochlamisus larva, and two possible food plants, red maple and willow. Photos by Scott Justis/BugGuide.net, Mary Keim, and John Tann.

Some populations of the leaf beetle Neochlamisus bebbianae eat red maple, and others eat willow; each type grows better on their native host plant. Hybrids between the two species are possible, and they don’t grow as rapidly when raised on either host. This might mean that ecology — adaptation to the different host plants — is creating reproductive isolation between the two forms of Neochlamisus. But it might also mean that the two forms are genetically incompatible.

Many species are separated by intrinsic genetic incompatibility. In these cases, hybrids have reduced fitness, or die outright, because the two species have evolved separately in such a way that mixed genomes cannot produce important proteins correctly. One example was recently found in two lines of the wildflower Arabidopsis thaliana — both lines had duplicate copies of an important gene, and in each line a different copy mutated into non-functionality, so some hybrids between the two lacked any functional copies [$-a].

To differentiate between this kind of genetic incompatibility and ecological isolation, coauthors Egan and Funk conducted not one but two generations of hybridization between maple and willow Neochlamisus populations. In the first (F1) generation, they bred parents from each host-specialized type; but in the second they performed a “backcross,” breeding the F1 hybrids with mates from one or the other of the parental populations.

This produced a population of backcrossed hybrids with 3/4 of their genes from one parental type, and 1/4 from the other. If intrinsic incompatibility separated the types, then these backcrossed hybrids would grow poorly no matter what their host plant. However, if adaptation to separate host plants isolates the types, then backcrossed hybrids would perform better on the host plant of the type with which they shared more genes. This is what Egan and Funk found — backcrossed hybrid larvae grew faster on maple if they shared more genes with maple-type Neochlamisus, and similarly for willow.

References

Bikard, D., Patel, D., Le Mette, C., Giorgi, V., Camilleri, C., Bennett, M., & Loudet, O. (2009). Divergent evolution of duplicate genes leads to genetic incompatibilities within A. thaliana Science, 323 (5914), 623-6 DOI: 10.1126/science.1165917

Egan, S., & Funk, D. (2009). Ecologically dependent postmating isolation between sympatric host forms of Neochlamisus bebbianae leaf beetles Proc. Nat. Acad. Sci. USA, 106 (46), 19426-31 DOI: 10.1073/pnas.0909424106

For yucca moths, does (flower) size matter?

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ResearchBlogging.orgIn a paper just released online at Molecuar Ecology ahead of publication, genetic tests on moth larvae provide the latest piece to the puzzle of why there are two kinds of Joshua tree — because the tree’s pollinators need to match its flowers [PDF].

I’ve written extensively about the interaction between Joshua tree and its pollinators. Like all yuccas, Joshua tree is pollinated only by yucca moths. Female yucca moths collect pollen in special mouthparts and deliberately apply it to a yucca flower after laying eggs inside it. When the eggs hatch, the moth larvae eat some of the seeds inside the developing fruit. Yuccas prevent their pollinators from laying too many eggs by selectively killing flowers too badly damaged by egg-laying [$-a].



TOP: The two forms of Joshua tree (western type on left, eastern on right). BOTTOM: Scaled comparison of moth body sizes and tree pistils. To lay eggs in a flower, moths must drill from near the top of the pistil to the positions marked by dotted lines. Photo by jby, Illustration from Smith et al.(2010), figure 1.

This last element of the interaction may have had significant consequences for Joshua trees’ evolutionary history. Joshua trees are pollinated by two different species of moths, which occur in different parts of the tree’s range: the larger Tegeticula synthetica in the west, and the smaller T. antithetica in the east. Joshua trees pollinated by the two different moth species are themselves different, both in their overall shape, and in the shape of their flowers’ pistils — specifically, the length of the route that a moth must drill to lay her eggs [PDF].

How does this difference in flower shape affect Joshua tree pollination? If a larger moth attempts to lay eggs in a smaller flower, it may be do more damage to the flower than the “native” pollinator would, triggering the tree to kill the flower. On the other hand, smaller T. antithetica might be able to lay eggs in a larger western-type flower without this risk. If this is the case, moths probably can’t pollinate western trees with eastern pollen, but they might be able to do the reverse.

Such one-way pollen transfer between the two Joshua tree types could produce a population genetic pattern called “chloroplast capture.” Joshua tree pollen doesn’t contain the full genetic code of the tree that produces it — it lacks the genes contained in the chloroplast, the cellular structure that conducts photosynthesis, because pollen grains typically don’t have chloroplasts. The DNA in the cellular nuclei of newly-formed seeds is a mixture of nuclear DNA (nucDNA) from a pollen grain and from one of their “maternal” parent’s ovules, but they get all their chloroplasts, and chloroplast DNA (cpDNA), from the ovule. If moths carry pollen from eastern trees to western trees, then the seeds produced would contain western cpDNA, but also some eastern nucDNA.


Asymmetric pollen transfer can lead to eastern-type trees with western-type chloroplasts. Figure 2 from Smith et al.(2010).

This is what we’ve found in Joshua tree populations near the region where the two tree types and their pollinators come into contact. At these sites, trees look like the eastern type (meaning they likely have eastern nucDNA, though we haven’t tested that yet) but have cpDNA that matches nearby populations of western-type trees [PDF].

The genetic pattern is only suggestive of one-way pollen transfer between the two Joshua tree types, though. We haven’t yet tracked the movement of moths directly, or estimated whether they actually are less successful when laying eggs on the wrong tree type. The newly-published study provides exactly these data. My colleague Chris Smith placed glue traps on Joshua tree flowers at the contact zone to estimate how often adult moths of each pollinator species visited each type of tree in the mixed population. Adult moths were more likely to be trapped on their “native” trees, though they did show up on the other type sometimes.


A yucca moth larva emerges from a Joshua tree fruit in the lab. Photo by jby.

Chris and I then collected fresh fruit from trees in the contact zone, and caught yucca moth larvae as they chewed their way out. Chris and another coauthor, Chris Drummond, then identified the species of each larva based on their genetics (the two pollinators look very similar at that stage) — and in our sample, the pattern of specificity was even stronger than that in the adults. The larger moth species, T. synthetica, never emerged from fruits of the small-flowered eastern trees. The vast majority of larvae of the smaller T. antithetica were also found inside their “native” tree’s fruit — but a handful did emerge from large-flowered western trees.

This mechanism could create the genetic pattern we see in Joshua tree populations. Larger T. synthetica doesn’t seem to lay eggs in (or pollinate) small-flowered eastern trees, but smaller T. antithetica can occasionally lay eggs in (and pollinate) large-flowered western trees. This should create asymmetric gene flow, with pollen moving from eastern trees to western trees, but not the reverse. The two Joshua tree types may not yet be reproductively isolated, separate species — but we won’t know for sure without looking at the plants’ nuclear DNA. As it happens, I’m working on that right now.

References

Godsoe, W.K.W., Yoder, J.B., Smith, C.I., & Pellmyr, O. (2008). Coevolution and divergence in the Joshua tree/yucca moth mutualism The American Naturalist, 171 (6), 816-823 DOI: 10.1086/587757

Marr, D., & Pellmyr, O. (2003). Effect of pollinator-inflicted ovule damage on floral abscission in the yucca-yucca moth mutualism: the role of mechanical and chemical factors Oecologia, 136 (2), 236-243 DOI: 10.1007/s00442-003-1279-3

Smith, C.I., Godsoe, W.K.W., Tank, S., Yoder, J.B., & Pellmyr, O. (2008). Distinguishing coevolution from covicariance in an obligate pollination mutualism: Asynchronous divergence in Joshua tree and its pollinators. Evolution, 62 (10), 2676-87 DOI: 10.1111/j.1558-5646.2008.00500.x

Smith, C.I., Drummond, C., Godsoe, W.K.W., Yoder, J.B., & Pellmyr, O. (2010). Host specificity and reproductive success of yucca moths (Tegeticula spp. Lepidoptera: Prodoxidae) mirror patterns of gene flow between host plant varieties of the Joshua tree (Yucca brevifolia: Agavaceae). Molecular Ecology DOI: 10.1111/j.1365-294X.2009.04428.x

Aphid-tending ants cull the sick from the herd

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ResearchBlogging.orgJust released online at Biology Letters: aphid-tending ants have been observed to selectively remove sick members of their “herd” [$-a].

Most aphid species produce some sort of sweet honeydew as waste while feeding on their host plants; ant-attended aphid species use this honeydew to attract ants. In many cases, the ants “milk” the aphids by stroking them to prompt release of the honeydew. While exploiting a colony of aphids, ants defend it as a food resource, protecting the aphids from predators. Aphid species that commonly rely on ant protection often lack defensive adaptations [$-a] found on species that don’t interact with ants.


Ants tend aphids on a milkweed plant. Photo by dmills727.

Niesen et al. report the results of experiments performed ants attending colonies of milkweed aphids, Aphis asclepiadis, which are susceptible to a fungal pathogen that can wipe out aphid colonies in a matter of days. In two experiments, they introduced aphids into the ant-attended colonies, and tracked what the ants did to them. They found that

  • Ants were more likely to remove the corpses of fungus-killed aphids than either the corpses of aphids killed by freezing or introduced live aphids; and
  • Ants were more likely to remove live aphids contaminated with fungal spores (conidia) than live aphids without spores.

The authors speculate that this behavior is a re-application of ants’ treatment of their own sick and dead within the colony. It seems clear that it should have benefits to both ants and aphids in this new context, slowing or preventing the spread of the fungus within an aphid colony. This benefit isn’t directly tested by Nielsen et al., but such an experiment is a logical next step.

Reference

Nielsen, C., Agrawal, A., & Hajek, A. (2009). Ants defend aphids against lethal disease Biology Letters DOI: 10.1098/rsbl.2009.0743

Way, M. (1963). Mutualism between ants and honeydew-producing Homoptera. Ann. Rev. Entomology, 8 (1), 307-44 DOI: 10.1146/annurev.en.08.010163.001515