Why does the eye care about the nose?

The ear, the nose, the eye: all of the neurons closest to the environment are doing on thing: attempting to represent the outside world as perfectly as possible. Total perfection is not possible – you can only only make the eye so large and only need to see so much detail in order live your life. But if you were to try to predict what the neurons in the retina or the ear are doing based on what could provide as much information as possible, you’d do a really good job. Once that information is in the nervous system, the neurons that receive this information can do whatever they want with it, processing it further or turning it directly into a command to blink or jump or just stare into space.

Even though this is the story that all of us neuroscientists get told, it’s not the full thing. Awhile back, I posted that the retina receives input from other places in the brain. That just seems weird from this perspective. If the retina is focused on extracting useful information about the visual world, why would it care about how the world smells?

One simple explanation might be that the neurons only want to code for surprising information. Maybe the nose can help out with that? After all, if something is predictable then it is useless; you already know about it! No need to waste precious bits. This seems to be what the purpose of certain feedback signals to the fly eye are for. A few recent papers have shown that neurons in the eye that respond to horizontal or vertical motion receive signals about how the animal is moving, so that when the animal moves to the left it should expect leftward motion in the horizontal cells – and so only respond to leftward motion that is above and beyond what the animal is causing. But again – what could this have to do with smells?

Let’s think for a second about some times when the olfactory system uses non-olfactory information. The olfactory system should be trying to represent the smell-world as well as it can, just like the visual system is trying to represent the image-world. But the olfactory system is directly modulated depending on the needs of an animal at any given moment. For instance, a hungry fly will release a peptide that modifies how much a set of neurons that respond to particular odors can signal the rest of the brain. In other words, how hungry an animal is determines how well it can smell something!

These two stories – how the eye interacts with the motion of the body, how the nose interacts with hunger – might give us a hint about what is happening. The sensory systems aren’t just trying to represent as much information about the world as possible, they are trying to represent as much information about useful stuff as possible. The classical view of sensory systems is a fundamentally static one, that they have evolved to take advantage of the consistencies in the world to provide relevant information as efficiently as possible*. But the world is a dynamic place, and the needs of an animal at one time are different from the needs of the animal at another.

Take an example from tadpoles. When the tadpole is in a very dim environment, it has a harder time separating dark objects from the background. The world just has less contrast (try turning down the brightness on your screen and reading this – you’ll get the idea). One way that these tadpoles control their ability to increase or decrease contrast is through a neuromodulator that changes the resting potential of a cell (how responsive it is to stimuli), but only over relatively long timescales. This is not fast adaptation but slow adaptation to the changing world. The end result of this is that tadpoles are better able to see moving objects – but presumably at the expense of being worse at seeing something else. That seems like a pretty direct way of going from a need for the animal to code certain visual information more efficiently to the act of doing it. The point is not that this is driven by a direct behavioral need of the animal – I have no idea if this is due to a desire to hunt or avoid objects or what-have-you. Instead, it’s an example of how an animal could control certain information if it wanted to.

This kind of behavioral gating does occur from retinal feedback. Male zebrafish use a combination of smell and sight when they decide how they want to interact with other zebrafish. Certain olfactory neurons that respond to a chemical involved in mating signal to neurons in the retina – making certain cells more or less responsive in the same way that tadpoles control the contrast of their world (above). It appears as if the olfactory information sends a signal to the eye that either gates or enhances the visual information – the stripe detection or what-have-you – that the little fishies use when they want to court another animal.

The sensory system is not perfect. It must make trade-offs about which information is important to keep and which can be thrown away, about how much of its limited bandwidth to spend on one signal or another. A lot of the structure comes naturally from evolution, representing a long-term learning of the structure of the world. But animals have needs that fluctuate over other timescales – and may require more computation than can be provided directly in the sensory area. How else would the eye know that it is time to mate?

What this doesn’t answer is why the modulation is happening here; why not downstream?

 

* This is a major simplification, obviously, and a lot of work has been done on adaptation, etc in the retina.

 

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The retina receives signals from all over the brain, and that is kind of weird

As a neuroscientist, when I think of the retina I am trained to think of a precise set of neurons that functions like a machine, grinding out the visual basis of the world and sending it on to the brain. It operates independently of the rest of the system with the only feedback coming from muscles that move the eye around and dilate the pupils. So when someone [Philipp Berens] casually mentioned to me that yes, the retina does in fact receive signals from the brain? Well, I was floored.

I suppose I should not have been surprised. In fruit flies, there has been a steady accumulation of evidence that the brain sends signals to the eye to get it ready to compensate for any movement the animal will make. Intuitively, that makes a lot of sense. If you are trying to make sense of the visual world, of course you would want to be able to compensate for any sudden changes that you already know about.

It turns out that there is a huge mass of feedback connections from the brain to the retina in birds and mammals, something termed the centrifugal visual system. And inputs are sent via this system from both visual areas and non-visual areas (olfactory, frontal, limbic, and so on). So imagine – your eye knows about what you are smelling. Why? In order to do what?

The answer, it turns out, is that we don’t know. It sends all sorts of neurotransmitters and neuromodulators. The list of peptides it sends are long (GnRH, NPY, FMRF, VIP, etc) as is the list of regions that send feedback to the retina. It seems as if which regions send feedback to the retina is very species-specific, suggesting something about the environment each animal is in. But why?

This is a post long on questions and short on answers. It is more a reminder that the nice, feedforward systems that we have simple explanations for are really complex, multimodal systems designed to create appropriate behaviors in appropriate circumstances. Also it is a reminder to myself about how little I know about the brain, and how mistaken I am about even the simplest things…

I would love someone more knowledgable than me to pipe up and tell me something functional about what these connections do?

References

Repérant J, Médina M, Ward R, Miceli D, Kenigfest NB, Rio JP, & Vesselkin NP (2007). The evolution of the centrifugal visual system of vertebrates. A cladistic analysis and new hypotheses. Brain research reviews, 53 (1), 161-97 PMID: 17059846

Vereczki, V. The centrifugal visual system of rat. Doctoral Thesis. PDF.

The eye(s)

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(Photo by artofdreaming)

Ed Yong has an absolutely brilliant article in National Geographic about the receptical for neuroscience’s favorite sensory system, vision:

But simple eyes should not be seen as just stepping-stones along a path toward greater complexity. Those that exist today are tailored to the needs of their users. A sea star’s eyes—one on the tip of each arm—can’t see color, fine detail, or fast-moving objects; they would send an eagle crashing into a tree. Then again, a sea star isn’t trying to spot and snag a running rabbit. It merely needs to spot coral reefs—huge, immobile chunks of landscape—so it can slowly amble home. Its eyes can do that; it has no need to evolve anything better. To stick an eagle’s eye on a sea star would be an exercise in ludicrous excess…

“Insects and crustaceans have become so successful despite their compound eyes, not because of them,” says Nilsson. “They would have done so much better with camera-type eyes. But evolution didn’t find that. Evolution isn’t clever.”

Eric Warrant, Nilsson’s next-door neighbor at Lund University, takes a more lenient view. “Insect eyes have a much faster temporal resolution,” he says. “Two flies will chase each other at enormous speed and see up to 300 flashes of light a second. We’re lucky to see 50.” A dragonfly’s eye gives it almost complete wraparound vision; our eyes do not. And the elephant hawk moth, which Warrant has studied intensely, has eyes so sensitive that it can still see colors by starlight. “In some ways we’re better, but in many ways, we’re worse,” Warrant says. “There’s no eye that does it all better.” 
Our camera eyes have their own problems. For example, our retinas are bizarrely built back to front. The photoreceptors sit behind a tangled web of neurons, which is like sticking a camera’s wires in front of its lens. The bundled nerve fibers also need to pass through a hole in the photoreceptor layer to reach the brain. That’s why we have a blind spot. There’s no benefit to these flaws; they’re just quirks of our evolutionary history.

It does make you wonder what it is like to be a bat, so to speak. Consider the scallop:

The mantle of the bay scallop (Argopecten irradians) is festooned with up to 100 brilliant blue eyes. Each contains a mirrored layer that acts as a focusing lens while doubling the chance of capturing incoming light.

What is it like to have access to 100 eyes? It is not so simple as just imagining that you could ‘see more’. Our eyes sense more than what we just see (so to speak). In human retinas, melanopsin isn’t used to form images but to help entrain circadian rhythms. These neurons send information to a different part of the brain (the suprachiasmatic nucleus) in a way that we can fundamentally feel in a different way. Now imagine those 100 eyes: are they all there for the same thing? Is the feeling of one the same as the feeling of another?

 

How many smells can you smell, audience-friendly edition

If you have a keen memory, you might recall that there was a little kerfuffle a couple months back concerning ‘how many smells can you discriminate‘. Here’s a layman’s introduction to the basis of the question, by It’s Okay To Be Smart.

And yes, I’m mostly posting this because it refers to this blog in the ‘sources’ section 😉

How words make color

Go check out this great interactive explanation of how words represent colors in English versus in Chinese.

Color words in Mandarin Color words in English

Interestingly, the most common color words in Chinese are for red, green, and blue while in English they are blue, green and pink!

[via FlowingData]

The sound of silence

Sensory neurons receive input from the outside world and send these signals on to the rest of the nervous system. This makes the concept of ‘silence’ fairly intriguing: what happens when there is very little sensory signal for the rest of the brain to process? It is well known that, after a while, no sensory stimulation means massive hallucinations. But quiet – brief periods of weak or no relevant stimulus – is different:

In the mid 20th century, epidemiologists discovered correlations between high blood pressure and chronic noise sources like highways and airports. Later research seemed to link noise to increased rates of sleep loss, heart disease, and tinnitus. (It’s this line of research that hatched the 1960s-era notion of “noise pollution,” a name that implicitly refashions transitory noises as toxic and long-lasting.)

Sound waves vibrate the bones of the ear, which transmit movement to the snail-shaped cochlea. The cochlea converts physical vibrations into electrical signals that the brain receives. The body reacts immediately and powerfully to these signals, even in the middle of deep sleep. Neurophysiological research suggests that noises first activate the amygdalae, clusters of neurons located in the temporal lobes of the brain, associated with memory formation and emotion. The activation prompts an immediate release of stress hormones like cortisol. [neuroecology: really? all noises?]

…He found that the impacts of music could be read directly in the bloodstream, via changes in blood pressure, carbon dioxide, and circulation in the brain. (Bernardi and his son are both amateur musicians, and they wanted to explore a shared interest.) “During almost all sorts of music, there was a physiological change compatible with a condition of arousal,” he explains…But the more striking finding appeared between musical tracks. Bernardi and his colleagues discovered that randomly inserted stretches of silence also had a drastic effect, but in the opposite direction. In fact, two-minute silent pauses proved far more relaxing than either “relaxing” music or a longer silence played before the experiment started.

In light of this, I found the experiences of a hermit who has been living alone since the 1980s fascinating:

He explained about the lack of eye contact. “I’m not used to seeing people’s faces,” he said. “There’s too much information there. Aren’t you aware of it? Too much, too fast.” (Note: he may have asperger’s)

“But you must have thought about things,” I said. “About your life, about the human condition.”

Chris became surprisingly introspective. “I did examine myself,” he said. “Solitude did increase my perception. But here’s the tricky thing—when I applied my increased perception to myself, I lost my identity. With no audience, no one to perform for, I was just there. There was no need to define myself; I became irrelevant. The moon was the minute hand, the seasons the hour hand. I didn’t even have a name. I never felt lonely. To put it romantically: I was completely free.”

…”What I miss most,” he eventually continued, “is somewhere between quiet and solitude. What I miss most is stillness.” He said he’d watched for years as a shelf mushroom grew on the trunk of a Douglas fir in his camp. I’d noticed the mushroom when I visited—it was enormous—and he asked me with evident concern if anyone had knocked it down. I assured him it was still there. In the height of summer, he said, he’d sometimes sneak down to the lake at night. “I’d stretch out in the water, float on my back, and look at the stars.”