Category Archives: Hymenoptera

Miles From Home: How Ants can Navigate Long Distances (and back!) to Forage

Ants are well known for their extraordinary ability to find food, and bring back enough to feed their vast social organisations. Being able to form relationships with other organisms that digest food, carry objects 50 times their weight and achieve great feats of communication and learning all help them forage… but how exactly do they find their way? Recent research from the University of Edinburgh has concluded the fascinating story of how the Formicidae navigate.

Plot a Course! Direction of Travel

Ants can decide on a direction for walking by using the position of the Sun in their visual field, as specialised cells in their compound eyes can detect the UV polarised light emitted by the Sun. Ants can maintain the correct course, whilst decoupling information where their body is an which direction they are travelling in. They also make use of visual landmarks (such as leaf litter), olfactory and tactile cues, and some species use the Earth’s magnetic field for navigation. According to the researchers at the University of Edinburgh, the ants construct a more sophisticated representation than they thought possible from the small size of their ganglia (brains), and can integrate information from different modalities (and from different areas of the brain) into the representation of direction.

How Far? Keeping track of Distance

Day-foraging ants, such as those in the genus Cataglyphis, are able to navigate exceptionally long distance (up to 200 metres and back!) by recording the distance they have travelled as well as the direction. An internal pedometer helps the ant remember the number of steps taken and this information is integrated with the ‘optical flow’ of objects moving around their visual field (which is an illusion- of course it is actually the ant that moves). Rather than each ant randomly roving away from the hive in search of food, the successful ‘pioneer’ must communicate the location to her sisters so they can make a sortie to the high quality patch of forage en masse…

neivamyrmex_army_ants_raiding_trail.jpg

Follow the Leader: Scent Trails

The long line of ants that you are bound to see in tropical forests are formed from scent trails that allow them to navigate back home, even if it is 200 metres away and in the dark! The ability to find the shortest route back is a crucial adaptation for avoiding desiccation in hot and arid environments. However, in army ant species, a group of foragers who become separated from the main marching column can turn back and form a circular ant mill, and run round constantly until they die of exhaustion! Ants have also been recorded to carry each other along a route, if an older and more experience forager notices that an internal nest worker (which are less familiar with the outdoor environment) is off the trail.

Final Word

So ants are able to backtrack to the location of their nest using their memories and the Sun as a reference point, and the way they operate is very similar to a self-driving car. This new research gives a unique insight into how brains of ants (and other insects) operate, and will inspire the next developments in robot system building to mimic their functioning, which would especially be useful for robots that need to navigate in forested areas. Modelling the neural circuits in the ant brain will also be useful to simply understanding more about the complex behaviours of the fascinating family of insects.

Further Reading

http://www.cell.com/current-biology/fulltext/S0960-9822(16)31466-X

http://jeb.biologists.org/content/209/1/26

http://science.sciencemag.org/content/353/6304/1155

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Bees: How do they Combat Disease?

A honeybee hive sick with disease can spell the end of a colony. Recent research has shown bees can use vaccinations and nurse one another to protect themselves from and prevent certain diseases. But how exactly do they do it?

Vaccinations

Honeybees live in huge colonies that co-operatively rear brood (developing eggs and larvae), so must find some way of protecting the next generation against disease. To do this, the workers actually naturally immunize their young against certain diseases which they might encounter. The findings of a recent paper, published in the journal PLOS Pathogens, finally reveal how the important immune-signal protein vitellogenin works to do this.

It was found foraging workers pick up and bring back contaminated pollen and nectar to the hive, and workers create royal jelly using it. The bacteria picked up from the environment persists in the jelly, and is then fed exclusively to the queen. The pathogens are digested in the queen’s gut and stored in the queen’s fat body (an organ similar to a liver). Fragments of the bacteria are then ‘carried’ by vitellogenin, taken via the blood to developing eggs inside the queen. These young are now immunized, all without taking a step outside their hexagonal brood cell.

Now that we know how bees immunize their young against infection, scientists can work on synthesizing a vaccine to prevent commercial bee colonies from becoming infected with disease- possibly aiding the fight against the crisis of colony collapse disorder (CCD).

But not all diseases in bees can be fought with immunity inherited from a parent. So how else do honeybees fight infection?

Nursing

Much like communist societies, honeybee hives divide up the hugely varied workload between different ‘castes’ of the colony (workers, drones and queen), and further divide workers into roles based on their size and age. Older, more experienced workers may be more likely to forage for the colony, and act as guard bees (they have actually been found to patrol the entrance the hive!). Younger, more naive workers however are usually more suited to nursing duties, which include feeding and tending to the queen and brood, as well as medical specialists which provide sick workers with anti-biotic laced honey.

A recent study, published in Behavioural Ecology and Sociobiology, gave nurse bees infected with a Nosema ceranae parasite a choice of honey from different plants. Bees with a higher level of infection tended to eat more sunflower honey, which contains the most antimicrobial activity. It also reduced the level of infection in the bees by 7%. A separate study suggests different honeys are effective against different diseases the bees may encounter. For example linden honey was better at fighter off a an infection of European foulbrood whereas sunflower honey was more effective against American foulbrood.

Nurse bees have other medical roles to reduce infection in a hive. For example, they can act as undertakers and remove the corpses of dead bees from the colony, dumping them far from the entrance. This behaviour is used to avoid spreading infections from pathogens and entomopathogenic fungi that proliferate on the bodies of dying insects,

In both of these incredible behaviours, bees can vaccinate and immunize their brood and sister workers by means of medicinal honey and food contaminated with bacteria. But where do these come from in the first place?

Natural Remedies

Floral nectar typically contains plant secondary compounds (those used for defence by the plant) which possess antimicrobial properties. This can be very useful to bees. Before the publishing of a recent study in PLoS One, we knew little more than the fact pollinators can reduce their parasite load by consuming nectar containing compounds such as nicotine.

This recent research has indicated parasitized bumblebees are taking advantage of these plant secondary metabolites in the wild, such as iridoid glycosides, and have a strong preference for visiting flowers that possess them. This quality of bees to self-medicate, by altering their foraging behaviour whilst parasitized, has massive implications for their ability to fight disease

Honeybees have other sources of medicine besides anti-microbial nectar. They have been found to collect resin from plants and incorporate it into their nests, which may help stop fungal parasities from colonizing their hive. (A study showed bees collect more of the resin when infected with fungal spores)

Final word

The fact that honeybees and bumblebees have evolved so many different ways in which to fight disease implies the risk that our wild pollinators face, as well as just how long they have been co-evolving alongside their assailing antagonists. Climate change and other drivers have recently made the problem of disease much worse, and research into this need to be rapid if we are to help our plighted pollinators.

Further Reading

Erler, S., Denner, A., Bobiş, O., Forsgren, E., & Moritz, R. F. (2014). Diversity of honey stores and their impact on pathogenic bacteria of the honeybee, Apis mellifera. Ecology and evolution, 4(20), 3960-3967.

Gherman, B. I., Denner, A., Bobiş, O., Dezmirean, D. S., Mărghitaş, L. A., Schlüns, H., … & Erler, S. (2014). Pathogen-associated self-medication behavior in the honeybee Apis mellifera. Behavioral Ecology and Sociobiology, 68(11), 1777-1784.

Richardson, L. L., Bowers, M. D., & Irwin, R. E. (2015). Nectar chemistry mediates the behavior of parasitized bees: consequences for plant fitness.Ecology.

Salmela, H., Amdam, G. V., & Freitak, D. (2015). Transfer of immunity from mother to offspring is mediated via egg-yolk protein vitellogenin. PLoS Pathog, 11(7), e1005015.

Simone-Finstrom, M. D., & Spivak, M. (2012). Increased resin collection after parasite challenge: a case of self-medication in honey bees. PLoS One,7(3), e34601.

Diseases in bumblebees and honeybees

All species of bumblebee and honeybee have associated diseases and parasites that impact on the health of populations. Emerging infectious diseases (EIDs) are those that pose a risk to human welfare (directly or indirectly) that affect ecosystem service production such as pollination of flowers or health of livestock. But what are these diseases and what are the factors that exacerbate them?

One commonly cited cause for colony collapse disorder (CCD) of the american honeybee (Apis mellifera) is the mite Varroa destructor. Varroa carries and transfers the viruses deformed wing virus (DWV) and acute bee paralysis virus (both implicated in CCD). Affliction with varroa mite also tends to weaken the immune system of honeybees. ‘Hygienic’ colonies of honeybees are able to remove the mites from brood cells and the workers groom themselves to remove the mite and disrupt it’s life cycle- this is a form of ‘resistance’ to the mite.

Other common parasites of honeybees include acarine tracheal mites, nosema spp (fungus that infest intestinal tracts), small hive beetle, wax moths and tropilaelaps (mites). Bacterial diseases include american foulbrood and european foulbrood and fungal diseases include chalkbrood and stonebrood. Honeybees are also susceptible to dysentery (inability to void faeces in flight) and viruses such as chronic and acute paralysis virus, kashmir bee virus, black queen cell virus, deformed wing virus and cloudy wing virus.

Researchers have found that two of these honeybee diseases (DWV and Nosema cerenae) are capable of infecting adult bumblebees. Further field work found that 11% of bumblebees were infected with DWV and 9% with N. cerenae, compared with honeybee infection rates of 35% and 7% respectively. The most likely explanation for the disease incidence in bumblebees is infection by honeybees, but bee-keepers can reduce the spread of disease by regular brood comb changes.  It is thought that ecological traits of these pollinating insects (e.g. overlapping geographic ranges, ecological niches and behaviours) promotes cross-species transmission of RNA viruses. Social behaviour and phylogenetic relatedness of social pollinators is thought to further facilitate transmission within and between hosts.

More recent evidence has suggested that commercial colonies bred for crop pollination and honey production can carry diseases (parasite infections and over 20 viruses) and be a threat to native species. Researchers found that 77% of imported bumblebee hives were contaminated with up to 5 different parasites. There is an urgent need for further research into the health of wild and imported bees and improvement in monitoring and management practices for honeybee and bumblebee colonies

Fürst, M. A., McMahon, D. P., Osborne, J. L., Paxton, R. J., & Brown, M. J. F. (2014). Disease associations between honeybees and bumblebees as a threat to wild pollinators. Nature, 506(7488), 364-366.

Manley, R., Boots, M., & Wilfert, L. (2015). Emerging viral disease risk to pollinating insects: ecological, evolutionary and anthropogenic factors. Journal of Applied Ecology.

Social Wasp Hierarchy: Who becomes Queen?

home-slide-yellowjackets

Social wasps have a caste system which separates individuals into reproductive males, infertile female ‘workers’ and a single reproductive queen. Towards the end of the summer, the existing queen runs out of stored sperm to fertilise eggs with so produces eggs that will eventually develop into fertile females. As each juvenile stage (egg, larva and pupa) is spent in a brood cell, the young queens that emerge must therefore compete to become the ultimate reproductive queen. But what factors determine which young queen dominates the hierarchy?

Eusocial species of wasps usually have their hierarchy determined by morphology of individuals. In the european wasp, Vespula vulgaris, the larvae that have been fed the most nutrients (which eventually becomes the largest reproductive adult female) will become the queen. The location of each cell is directly related to the amount of food a larva can receive, so the queen cells are usually located at the bottom of the nest which encounters most of the foragers. Several candidates of queens arise, which then compete to create a hierarchy of queens for an ultimate queen to be selected. The precise reasons behind the variations in queens is unknown, but it is thought to be related to fat stores which elevate a queen’s quality.

However not all social wasps have castes with such a variation in size and structure of individuals. In polistine paper wasps and stenogastrines (hover wasps), the hierarchy of females is determined behaviourally through dominance interactions. These hover wasps do not have predetermined or rigid castes, and young females need to constantly assert dominance to climb a strict age-based inheritance queue to become the reproductive (queen). Paper wasp colonies are founded by multiple reproductive females, and one of these foundresses will acquire dominance over the others and become the sole reproducer, with the others becoming ‘helper females’.

All female wasps are potentially capable of becoming the colony’s queen, which is usually achieved by a wasp laying eggs first and constructing the nest. Multiple young females usually compete with each other by eating the eggs of rival females. The queen may simply be the female that eats the largest number of eggs whilst safeguarding her own (and laying the most). Once the eggs have hatched, the subordinate females stop laying eggs and instead forage for the new queen and feed her young. If the dominant female dies, a new hierarchy may be established with a former worker acting as the replacement queen.

Different genera of wasps have different strategies for deciding which female becomes the egg laying queen that will give rise to a new progeny. However all strategies end up with the strongest and most reproductively capable female becoming queen. One or a multitude of factors may influence how competitive a young queen is, which include morphology (size and structure), nutrition (fat stores) and behaviour (e.g. aggression). Paper wasps have been found to recognise individual faces, so more complex forms of communication (using facial cues) may be used in the competition to become queen.

How else do pollinators benefit from plants?

bees

Plants primarily attract pollinators by offering nectar rewards (as a source of sugar for energy) and pollen (as a protein source for developing young). In return, the pollinator will pass on the plant’s genetic material for sexual reproduction. Using this animal vector is much more efficient than relying on wind pollination, but how does the pollinator benefit from this interaction? Besides nectar and pollen, what other rewards can flowers offer? How can plants sometimes ‘deceive’ pollinators by providing a false reward?

Some neotropical plants, including orchids, produce scent which is collected by the males of Euglossine bees to use as pheromones for courtship. The bees secrete saliva full of lipids onto the floral surface, which absorbs the scent compounds and is then collected by the bee’s ‘corbicula’, a modified pollen basket. Whilst the bees are collecting these female-attracting scents, the orchid’s specialized anthers deposit a ‘pollonium’ onto the back of the bee.

Pollinators have also been found to consume floral tissues (the plant itself!) as a reward for assisting with reproduction. Beetle species often consume floral tissues but also act as pollinators. For example cycads are often pollinated by specialized weevils that eat the cycad ‘flowers’ called cones. To limit the damage done, the plants often produce low concentrations of secondary metabolite (toxins) that accumulate in the insect and this limits the amount of floral tissue the pollinator can eat.

Other specialized plant-pollinator mutualisms has the plant producing oils which are used by bees to build nests and feed larvae. In many cases these fatty acid secretions are made rather than nectar, which means a more specialized pollinator can co-evolve with the plant. This leads to more efficient pollination as the insect is limited to only travelling and distributing pollen between a few plant species.

Sometimes pollination occurs by deceit. The dead horse arum produces chemical compounds that mimic those produced by carcasses. This attracts carrion flies and traps them in the flower overnight, covering them with pollen. Orchids of the genus Ophrys emit compounds which mimic female wasp pheromones and have visual displays that look similar to female wasps. The male wasp visits and attempts to mate with the flower, but inadvertetly pollinates it if fooled twice! Neither of these plants reward the pollinator with food (or otherwise) for pollinating it- this one sided ecological interaction is known as ‘commensalism’.

Many different incentives to attract insect pollinators to specialise on a particular species (or genus) of plant exist. The bizarre strategy of the fig tree has a chalcid wasp insert its long ovipositor to lay its eggs inside the fig fruits. The fig tree grows it’s flowers inside the fig fruits so the wasp actually pollinates it in return for the plant providing a habitat and food source for the developing larvae. Other plants have been shown to ‘cheat’ and deceive their pollinators into accepting a non-existent reward (like the wild orchid wasp mimic). However recent research has found that bumblebees have a preference for plant scent compounds that give an honest indication of reward quality. Plant resources put towards reward quality is limiting however, so an equilibrium exists between the evolution of cheaters and ‘honest’ signallers.

 

Irwin, R. E., Adler, L. S., & Brody, A. K. (2004). The dual role of floral traits: pollinator attraction and plant defense. Ecology, 85(6), 1503-1511.

Knauer, A. C., Schiestl, F. P. (2014), Bees use honest floral signals as indicators of reward when visiting flowers. Ecology Letters. doi: 10.1111/ele.12386

Simpson, B. B., & Neff, J. L. (1981). Floral rewards: alternatives to pollen and nectar. Annals of the Missouri Botanical Garden, 301-322.

How do Social Insects make Decisions?

beesmed

Using information passed on by others can greatly improve individual fitness, and has been the fundamental mechanism underlying the evolution of social insects such as bees, wasps, ants and termites. However in some situations it is better to ignore social information and for an individual to use its own prior knowledge and experience. So how do these colony-forming insects tailor their reliance on social information for the benefit of the ‘superorganism’? Scientists have recently reviewed the literature and made theories as to the nature of decision making in insects.

Social information is relatively ‘cheap’ to obtain for hymenopteran foragers, because they can bypass the costs associated with exploration and food sources obtained socially are likely to be better quality. In the truly eusocial western honeybee, Apis mellifera, generations overlap so information passed on by the ‘waggle dance’ (movements conveying location and quality of food sources) increases the fitness of that colony. Foraging choice are further refined by chemical cues (pheromone trails) and simply presence of other foragers.

Relying on social information may also incur costs and may not lend an evolutionary advantage. In the case of the ant forager, if she ignores social information she may find a novel food source that will benefit the colony as a whole, whilst a well-used food source is depleted (I.e. exploration produces more up-to-date information). Honeybees that rely on dance information may take time to find a dancer and may need multiple viewing and excursions to find the communicated food source.

A trade-off between these advantages and disadvantages will adjust how often (and what proportion of) social insects rely on social information. All animals tend to display the most profitable information they know, so relying on social information may be more profitable than exploration. For example honeybees only communicate their dance after finding high quality food sources. ‘Social learning strategies’ in animals are genetically determined in response to environmental and social cues. One such approach is the ‘copy if dissatisfied’ strategy, where animals will use social information if their current information is below a fitness ‘threshold’. These optimum social learning strategies can also be acquired (ironically) through social learning.

Grüter, C., & Leadbeater, E. (2014). Insights from insects about adaptive social information use. Trends in ecology & evolution, 29(3), 177-184.