Showing posts with label gustatory receptor. Show all posts
Showing posts with label gustatory receptor. Show all posts

Wednesday, March 5, 2014

Taste And Be Tasted – Fair Is Fair

Biology concepts – metamerism, tagmentizaton, taste, arthropods, receptor, parasitism


Carbonated sodas come in all flavors, but across all cultures, it is
the carbonation that is the same. Mauby is a tree bark flavored
soda sold in the West Indies islands of Bermuda, Trinidad and the
like. On the left is a bird’s nest/white fungus soda sold in Vietnam.
I don’t think it includes the bird.
There is no doubt that humans love the taste of carbonated sodas. There are as many flavors as you can imagine, but the common element among them is the infusion of carbon dioxide (CO2). Do you taste the carbon dioxide or is it important for some other reason?

Many insects will tell you that it’s the CO2 that makes the difference. Fruit flies, mosquitoes, ticks and other insects can taste CO2 on surfaces and in the air. For mosquitoes and ticks, tasting CO2 helps them find food. These are hematophagous (blood-eating) organisms, and they find their victims by flying upstream along their exhaled CO2 and the CO2 that is exuded from their skin.

Even more amazing, fruit flies and other insects taste the increased CO2 that stressed (injured, diseased) flies emit. They may avoid other insects that are dying so they won’t be near disease or danger. In other insects, they may follow it to animal carcasses - their buffets. In either case, the insects can actually taste death.

But are they tasting CO2? It’s a gas, and we have said that gases are detected and perceived by smell, not taste (except for us and DMS). It turns out that CO2 sensation is really an exception. A 2007 paper from John Carlson’s group showed that the receptor heterodimer (hetero = different, and dimer = two different proteins) is made of GR21a and GR63a, two gustatory proteins (hence the GR in the name).

However, the two taste receptors are located on olfactory neurons. The signal is detected by taste signaling on a smell neuron, and the signals are then sent to the smell portion of the brain! This may be one of the biggest exceptions in all of taste science, and it’s the insects that have it and use it.

For insects to accomplish many different tasks with taste, it helps to have the taste receptors in specific places. Catfish had them all over their body, but that’s not very specific. In insects they are found in distinct places, and may have distinct functions.


The shrimp is a good example to show metamerism in arthropods.
All the parts are just reiterations of the same subunit. Some kept
their appendages, and some changed them into something else.
Tagmentization is the result of modifications so that some of the
somites act together as the cephalothorax, and others
from the abdomen.
Many arthropods have taste receptor sensilla on exterior mouthparts, on their legs, on their antennae, and even on their wings. These may seem like a lot of work to develop them on so many different structures, but maybe not. Metamerism is at work.

Metamerism (meta = subsequent, and mer = unit) is a biology concept for efficient addition of complexity in an animal. Over time and evolution, certain specific structures and functions may develop in response to pressures. It is much more efficient to just create another unit using the same blue prints instead of creating a new part from scratch. The repeat is metamerism; the specialization over time of the different mers is called tagmatization.

You can see metamerism and tagmentization at work in arthropods and annelids (worms) by looking for repeating units. Millipedes and centipedes are great examples. Their bodies are made from many copies of the same basic unit. In many animals, repetition of units allows for drift over time and slow changes in structure and function, even grouping of different mers together for special function (tagmatization).

Mers (or somites) in insects include appendages like legs. But over time, many of the appendages evolved into other structures, like mouthparts, antennae, and egg-laying apparatus. Some characteristics are retained, others are dropped or altered, and some new characteristics appear.


Feel like your being stared at? The left picture is good for showing
the mouthparts of a grasshopper. Every one is a remnant of an
appendage. The mostly come in pairs, one from each modified
appendage. On the right, the cartoon shows the different
mouthparts, the labrum (lr) and hypopharynx/labium (hp/lb)
have fused to form just one piece. md = mandible, mx = maxilla
In terms of taste, the appendages seem to have been a seat of gustatory receptor sensilla. When several appendages evolved into mouthparts, the taste receptors were there. When some appendages developed in antennae, the taste receptors were there. But there is still the chicken and the egg question - did taste receptors on mouthparts result from them being derived from appendages, or did taste receptors on legs and other appendages come from early appendages being used as mouthparts?

A run down of tasting anatomy is hard for insects as a whole, because different arthropods taste with different parts, but some structures are more common. Mouthparts seem to be a favorite, and that makes sense. Flies taste with their probsocises (am I making up the plural?), but they also taste with the ends of their legs. Arthropod legs come in segments, and the last segments are called the tarsi.

Flies can taste food with their tarsi just by landing on it, but the also have taste receptors higher on their legs as well. Honeybees taste primarily with their antennae, but other flying insects can actually taste things with their wings! Wing tasters include fruit flies and mosquitoes, and they are more of an exception than you might think. We talked above about how tasting with different parts isn’t so crazy, since metamerism is just the modification of similar starting parts. But wings are not modified appendages.

Wings actually evolved from abdominal gills, and most insects have either given up these early structures and those that have them don’t taste with them. It may be that taste receptors on wings developed on their own, or that taste is older than metamerism. We don’t know their function yet – you work on that one.


Drosophila is the quintessential research model. The left
cartoon shows the olfactory and gustatory receptors. Notice
how many taste receptors are around the proboscis. On the
right, the red dots show all the different places gustatory
receptors are found. Wing margins, legs, tarsi, and mouthparts
all have taste receptors.
We have introduced mosquitoes and taste when we talked about CO2 above, but they come into play again here, according to a 2010 study. They taste with wings, and this may have something to do with how we can keep mosquitoes away from us. The two main chemical deterrents to mosquitoes are DEET and citronella candles. And they work differently.

Citronellal is only smelled by mosquitoes; the active molecule triggers only olfactory receptors. But DEET triggers both olfactory and gustatory receptors, it is smelled and tasted. Both senses stimulate avoidance responses in insects, so even if a mosquito lands on you, the DEET you put on will be tasted and may keep it from biting.

So some insects taste with wings - is that as weird as it gets? Nope, some females taste with their ovipositors (ovi = egg, and posit = laying). Ovipositors are a result of metamerism, they are modified appendages. The females of many species can taste the plants or places they land to determine if they are a suitable place to lay eggs.

The ovipositors most likely have rare taste receptors, applied to only to this one specific task. For example, there are two subspecies of a particular fruit fly called a goldenrod gall fly (Eurosta solidaginis). The females look for specific plants, and then for buds of the right age in which to insert their eggs. The growing larvae then feed on the bud, and cause a tumor (gall) to form.


The ovipositor of a female wasp or fruit fly is also a modified
appendage. In the wasp on top has a rigid ovipositor that may
be used to inject eggs into a caterpillar larva. On the fruit fly
ovipositor below, you can almost see the sensillae that
contain the taste receptors.
The interesting point is that there are two different kinds of goldenrod and two different kinds of flies. One type of fly will never pick the other type of goldenrod to lay it eggs on. The slightly different plants must have slightly different tastes, and the two subspecies of flies have evolved to react favorably to only one of the two tastes.

Obviously, some insects pick their plants very carefully. Let me give you an example that really knocks this point home. Tiger moth (Grammia incorrupta) caterpillars are sometimes parasitized by flies or wasps that lay their eggs inside the wooly bear (tiger moth caterpillar). A 2009 paper shows that when this occurs, the caterpillars switch the kind of plant food they eat, opting for poisonous plants that contain pyrrolizidine alkaloids (PA).

The PA-rich food is much less nutritious than the caterpillar’s regular food, so it definitely costs the caterpillar in terms of grown and health, but the PA is toxic to the parasites. The food choice sometimes depends on the number of parasitic eggs laid in one individual caterpillar. Just one egg – a caterpillar may eat some PA-rich plant material and let its immune system do the rest of the work. But with more eggs, the woolly bear will consume PA-rich plants exclusively – hoping to kill off all the eggs. The caterpillars are self-medicating, tasting their way back to health.

Turnabout is fair play – we haven’t discussed the plants that are being eaten by all these insects. In some cases, it turns out that the plants are tasting them right back, and even tasting each others' messages.


You can see the parasitic wasp injecting eggs into the
caterpillar. When the eggs hatch, they will feed on the
caterpillar through their larval stage. Two things may
happen. The caterpillar may switch plants (based on taste)
to try and poison the parasites. Second, the plant they are on
now may have called in the wasps to kill the caterpillar using
volatile chemicals, and the toxic plant that the caterpillar
switches to may do it again.
Corn plants (maize) get munched on by caterpillars. In response, they produce chemicals to attract predators of the caterpillars. This has been known for a while. But a 2000 study showed that the plants respond to the caterpillars saliva; the maize tastes it (contact chemosensation) and starts to send out the volatile chemicals that will attract parasitic wasps looking to lay eggs in the caterpillars. A more recent study shows that the caterpillars play an even bigger role in their own demise.

The volatile chemical that maize uses comes in two forms; it’s the switch from primarily one form to the other that attracts the wasps. But even before the plant starts to produce the attractive form, the caterpillar’s saliva converts the inactive form to the attractive form. The attractive message starts about a day before the plant starts to make the attractive form. The maize molecule has evolved to make the caterpillar call the cops on itself.

What is more, plants can send taste messages to nearby plants through the dirt. In a 2011 study, researchers induced drought like conditions on one row of plants. In less than an hour, plants five rows away started to close their stomata (pores in leaves) to conserve water for an impending drought. Plants that were just as close, but planted in a different container did not prepare for drought, so the message had to be traveling through the soil. I leave it to you to decide if this is really a taste sense.
           
So - if you’re a raw food enthusiast, you might be being tasted back. And maybe your food is spreading the word about you to his neighbors. Next week – why do we call spicy food "hot?"



Falik O, Mordoch Y, Quansah L, Fait A, Novoplansky A (2011). Rumor Has It…: Relay Communication of Stress Cues in Plants. PLoS ONE, 6 (11)

Lee Y, Kim SH, & Montell C (2010). Avoiding DEET through insect gustatory receptors. Neuron, 67 (4), 555-61 PMID: 20797533

Singer, M., Mace, K., & Bernays, E. (2009). Self-Medication as Adaptive Plasticity: Increased Ingestion of Plant Toxins by Parasitized Caterpillars PLoS ONE, 4 (3) DOI: 10.1371/journal.pone.0004796

Allmann S, & Baldwin IT (2010). Insects betray themselves in nature to predators by rapid isomerization of green leaf volatiles. Science (New York, N.Y.), 329 (5995), 1075-8 PMID: 20798319

Kwon JY, Dahanukar A, Weiss LA, & Carlson JR (2007). The molecular basis of CO2 reception in Drosophila. Proceedings of the National Academy of Sciences of the United States of America, 104 (9), 3574-8 PMID: 17360684



For more information or classroom activities, see:

Carbon dioxide taste in insects –

Parasitic wasps –

DEET/citronella –

Plant volatile defense chemicals -



Wednesday, February 26, 2014

Strange Insect Tastes

Biology concepts – sensilla, uniporous sensilla, gustatory receptor, RNA world hypothesis


Chalcanthite (copper containing) is used in traditional
medicines, even though it is toxic. Recent study shows
it is anti-inflammatory when mixed with egg white. A
2013 study shows that after burning off the water from
chalcanthite and mixing it with egg white to eliminate
toxicity, the concoction inhibits several signaling
pathways that would otherwise stimulate
inflammation. I’m not sure you would have to taste
this mineral to know what it was.
Yukon Cornelius, you know- the prospector of Rudolph the rode-nosed reindeer fame, used to taste the end of his pick after ho tossed it into the air and it stuck in the ground. He was looking for gold, and the action implied that he could know if there was gold in the ground by how it tasted.

Don’t laugh, I’ve seen many a geologist taste a rock to get some information about it and the area from which it came. Halite is an easy one (Greek hals = salt), but other minerals have tastes as well.  

Borax is sweet and alkaline. Ulexite tastes alkaline while chalcanthite is sweet and metallic (see picture). There are others as well, just be careful that you have a clean area to taste, not one that has been exposed to overflying birds.

Believe it or not, this does have application in biology. Geologists are looking for older and older borates, like borax and ulexite, because there are hypotheses that these might have been important in stabilizing the ribose of RNA in the prebiotic RNA World. They field test samples by tasting to find borates to test for age.

But I don’t know about Mr. Cornelius using taste to find gold. Gold is famously inert, it shouldn’t have any taste at all. In fact, professional ice cream testers use gold spoons just so they won’t be influenced by anything but the product.

It’s not probable, but perhaps geologists and ice cream tasters learned from insects about taste. Arthropods are famous for finding interesting uses for taste. They taste their way to food, to find mates, and even to find appropriate places to lay eggs. But what is most exceptional is how and where they do it. Insects are our big exceptions for the day.

Insects (just one class of arthropods) are invertebrates, so they don’t have the traditional taste buds associated with mammals, fish, and birds. Instead they have taste receptors house inside gustatory sensilla. There are several types of sensilla on insect bodies, and different types can house different senses. Gustatory sensilla are usually of the trichome (or trichoid, meaning three faces) type, meaning that they look like hairs and most people call them hairs. They are not hairs because they are not made of keratin protein. They are made from chitin, the same material as the crunchy insect exoskeleton, and they are hollow.


This is a photomicrograph of an insect antenna, with
several different types of sensilla. Look closely to see
the differences. The sharper ones come in two lengths,
shorter (b) and longer (a), but they are both considered
trichome type. Others are blunter and thicker, these are
the basiconic type (c). Trichoid house touch and taste
receptors. Basiconic hold olfactory (smell) receptors.
The end of the sensilla has a single opening, a very small opening (10 nm/4 x10-7 in). Because of the single opening, it is also called a uniporous sensilla.  The pore is so small that water molecules trapped within it can’t evaporate. If the sensilla then rubs up against a surface and some surface molecules get trapped in the water. These molecules can stimulate the gustatory receptors and the insect can taste what they have touched.

These sensilla are common among the arthropods. The arthropod phylum includes the chelicerae (scorpions, spiders, etc), myriapoda (millipedes and centipedes), hexapods (insects, beetles, ants, butterflies, etc.), and crustaceans (crabs, lobsters, shrimp, etc.). I was surprised to find that crayfish and lobsters have sensilla on their legs – antennae, O.K., but on those hard legs? Spiders, even the ones that aren’t hairy, have gustatory and olfactory sensilla, as do millipedes and similar. Since there are more than 500 arthropod species for every mammalian species on Earth, it would seem that uniporous sensilla are the common way to taste. Taste buds like ours are the exception.

The gustatory receptors of arthropods are located on neurons housed within the sensilla. The neuron depolarizes based on the strength of sense signal and transmits that information to the central nervous system. We’re talking about insects here, so “brain” isn’t really the right word to use. In arthropods, both gustatory and olfactory inputs go to an olfactory center, so one could argue if it is really a taste they are sensing. It’s perceived in the smell center, but comes from direct contact, not a gaseous molecule from a distance – let the discussions begin.

However they perceive it, gustatory receptors in uniporous sensilla of insects are used for a variety of purposes. That’s not exceptional – we do too. But insects tend to do it better and for more varied reasons. The obvious function is for finding food, but here insects take shortcuts that other animals do not. Take, for instance, one of the most important research animals ever, the fruit fly (Drosophila melanogaster).


Top left: the proboscis of the fruit fly can be seen projecting
forward, like an insect elephant trunk. Top right: the
proboscis monkey, while at the bottom is the Darwin’s hawk
moth. The term “proboscis” can be used with invertebrates
or vertebrates, but it means different things in each case.
When speaking invertebrate, proboscis means tubular
mouth parts. In the vertebrate tongue, it doesn’t mean
tongue at all, it means nose or snout.
The fruit fly shortcut is seen when it tastes something sweet or tastes water, the neural circuitry automatically causes muscular movements that extend the fly’s proboscis (pro = forward, and boskein = to feed) for feeding. Similarly, if the fly tastes something bitter, the reflex is for the proboscis to retract and feeding to either stop or not start. Muscle movements are connected directly to taste sensation. We can’t do that, unless you count the face kids make when they eat broccoli.

Also in feeding, it seems that insects are the exception to the rule of a limited number of taste receptors. Insects that feed on only one type of plant (feeding specificity) tend to evolve taste receptors for chemicals made only by that particular plant. This leads to many receptors with novel and new specificities.

Depending on the diet or other uses for receptors, insect species also vary greatly in the number of different receptor genes they have. Fruit flies have more than 70 different receptor genes, while honeybees may have as few as ten. However, they probably all have a receptor for tasting water.

The water receptor was recently identified in Drosophila. A 2010 paper in Nature describes a gene and protein called pickpocket 28 (ppk28). The scientists tracked the brain responses to water in flies when the gene was stimulated or when they removed the gene.  There was more activity when plain water was given than when it was mixed with salt or sugar (more water if no solute). When mutated to become nonfunctional, flies drank water for much less time (3 seconds versus 10 seconds) and showed much less brain activity in response to water.

Studies like this are hard, but the amount of brain space devoted to sensing things like this makes it easier. Undoubtedly, the ants (still in the arthropod phylum) are king when it comes to sensing taste and smell. For example, worker ants can distinguish between different single and double sugars, meaning that they differentiate many subtle differences in taste.


Here we witness a heartwarming act from the animal world.
C. japonicus ants have adopted the fusca caterpillar out of the
goodness of their hearts. Don’t even consider that the
caterpillar has tricked the others into think he is an ant, or
that the ants feed him just so he will secrete sugars that the
ants can snack on. No way - it’s a Hallmark special, not a
reality TV episode.
But they can also select different sugars based on immediate needs; whether they would be good for stored energy or immediate energy. This suggests nutrient sensing as well as taste differentiation. For example, Camponotus japonicus ants have a symbiotic relationship with Niphanda fusca butterfly larvae (caterpillar).

The ants protect the caterpillars because the caterpillars provide the ants with sugars, specifically, a disaccharide called trehalose + an amino acid, glycine. The ants actually adopt the caterpillars and raise them with their colony because the caterpillars secrete a pheromone that mimics that of the ants. The ants accept them because they taste like one of their own. The gift of sugar reinforces the relationship.

Other ant species will select different sugars equally, but the C. japonicus ants prefer trehalose. What is more, those ants prefer trehalose + glycine even though no ant species prefers glycine alone. So in this species, symbiosis drove evolution of a taste receptor for trehalose, and is modified by glycine.

One last example on ants, leaf cutter ants (Atta vollenweideri) produce different taste receptors based on the caste they belong to (large workers, small worker, soldiers, queens). A 2013 paper shows that the different castes and subcastes also have different numbers of taste receptors on their legs and antennae, so they respond to stimuli differently. One stimulus might be cohorts (members of the same colony) or, in other insect cases, for finding mates.

Drosophila fruit flies have been studied for this as well. A 2012 paper showed that different gustatory receptors and different pheromones are found on male and female flies. Fruit flies perform many different courtship rituals, and these take energy and time. It wouldn't pay to be courting another male – so sensing whether another fly is male or female is important.


Courtship in fruit flies is more complex than teenage dating.
The motions shown above are just a few of the actions that
take place, but the others aren’t G rated, so I decided not to
show them. The male doesn’t pick the female based on looks,
smarts, or ability to stick to a budget. New research shows
that he picks her based on the fact that she doesn’t
taste like a male.
This study mutated certain taste receptors. When male receptors are stimulated by male pheromones, courtship rituals stop. But if a female is tasted, the courtship continues. In the mutant flies, males would court males. When only mutant males were placed together in a container, they would line up in a long line, one fly courting the male ahead of it and being courted by the male behind it.
In butterflies, these different receptors give clues about evolution.  In the postman butterfly (Heliconius melpomene), researchers found 73 putative gustatory receptor genes, but the number of copies of gene and the variations of some genes varied between males and females. Fully one third of the genes shows a female bias in expression level, many being found on female legs, but not male legs. The results also showed that many of these were also the result of many recent gene duplications. 
Gene duplications allow for more genetic drift, and this would result in a greater number of possible receptors. Varied expression suggests that females are using the receptors for things that the males are not. So female behaviors seem to driving the expression and evolution of the gustatory receptors in butterflies. Once again, the women are in charge.

Next week, we should look at the weird places insects have taste receptors and how taste plays a role in egg laying. Even weirder, insects may taste plants, but it turns out that the plants are tasting them right back.



Choi EA, Park HY, Yoo HS, & Choi YH (2013). Anti-inflammatory effects of egg white combined with chalcanthite in lipopolysaccharide-stimulated BV2 microglia through the inhibition of NF-κB, MAPK and PI3K/Akt signaling pathways. International journal of molecular medicine, 31 (1), 154-62 PMID: 23128312

Briscoe AD, Macias-Muñoz A, Kozak KM, Walters JR, Yuan F, Jamie GA, Martin SH, Dasmahapatra KK, Ferguson LC, Mallet J, Jacquin-Joly E, & Jiggins CD (2013). Female behaviour drives expression and evolution of gustatory receptors in butterflies. PLoS genetics, 9 (7) PMID: 23950722

Koch SI, Groh K, Vogel H, Hansson BS, Kleineidam CJ, & Grosse-Wilde E (2013). Caste-specific expression patterns of immune response and chemosensory related genes in the leaf-cutting ant, Atta vollenweideri. PloS one, 8 (11) PMID: 24260580

Thistle R, Cameron P, Ghorayshi A, Dennison L, & Scott K (2012). Contact chemoreceptors mediate male-male repulsion and male-female attraction during Drosophila courtship. Cell, 149 (5), 1140-51 PMID: 22632976


For more information or classroom activities, see:

Mineral taste –

Uniporous sensilla –

Proboscis –

Niphanda fusca

Leaf cutter ants -