Showing posts with label form follows function. Show all posts
Showing posts with label form follows function. Show all posts

Form Follows Function - It’s About Time

Biology concepts – circadian rhythm, vision sense, adaptation, parasitism, form follows function


The sun and the moon are symbols of different
activity cycles. As with everything else, we have to give
them human characteristics (anthropomorphism).
Many animals are active in the day or the night, but not both. So what are humans, diurnal (active in the daytime), nocturnal (active in the nighttime), or something else?

Maybe humans are two species, because I know folks who can’t accomplish anything before noon, and do their best work after 11:00 pm, whereas I get up around 5:00 am and am pretty much useless after 8:00 pm.

Whether diurnal or nocturnal, organisms are physically and behaviorally adapted to their activity pattern. This includes the way they sense their environments. Diurnal animals are more likely to have color vision, while nocturnal animals may only see in black and white. The upside for nocturnal animals is greater visual sensitivity, so they can see better than diurnal animals in low light conditions.

The reasons for these different visual talents lies in the types of light receptors on the retina. Rods sense light, but only its presence or absence (white/black). Different receptors, called cones, detect various wavelengths of light (colors). Diurnal animals have about 5-10 times more cones than nocturnal animals (3 types, one for yellow, one for green to violet, and one for red to orange), but they only function in higher levels of light. Therefore, the greater number of rods in nocturnal animals allow for more sensitive night vision, a good thing to have if you are active after sundown.


Rods (yellowish) and cones (blue) are different light receptors located on the retina. Rods are more numerous and detect low levels of light. Cones are less numerous and sense colors of light, but require more light. As shown in the middle image, the tapetum is located beneath the retina in some animals, and can bounce light back to the retina. This bouncing around is responsible for animals glowing eyes at night.
Many nocturnal species have an additional adaptation to improve their night vision. Their retina has an iridescent layer called the tapetum lucidum that bounces the available light around so it may hit more rods. This improves sensitivity, but at a cost to acuity (the image gets a little fuzzier). When you shine a flashlight in the woods at night, the little pairs of reflections you see are the tapetum lucida of the animals looking back at you. The light bounces around inside the eye and some escapes back out through their pupils and that is what you see. Some look at your flashlight to see if you are a predator, others look to see if you are worth eating.

But not every animal with a tapetum lucidem is necessarily nocturnal. An interesting new study has looked at the visual system of the Peter’s elephant nose fish (Gnathonemus petersii). This weakly electric fish has a long nose-like appendage that was thought to mediate location and communication through electrical pulses. But scientists at the University of Cambridge have found that this fish has surprisingly good vision to go along with electrical impulse usage.

The elephant nose fish lives in the dark, murky waters of Central Africa. For this low light environment, it has evolved a unique retinal arrangement for its rods and cones. The cones are arranged in discrete packets, each housed in a cup lined with a tapetum lucidem. Behind these cones are the rods that work in lower level light. In this way, the visual field can respond with cones and rods at the same time. It is believed that this gives the elephant nose fish the ability to pick out predators moving quickly through its visual field.
 
Humans don’t have a tapetum lucidum, so when reflected light bounces off our retinas and back out the pupils, they appear red like the retinal blood vessels and tissues. This is the eerie red eye effect on some flash photography. I always thought it was a sign of vampirism!

Other nocturnal animals, like many owls, rely on hearing and smell more than vision. They are adapted to maximize these senses. We have discussed previously the changes in owl anatomy (Do You Have To Be Ugly To Hear Well) as examples of form following function to improve hearing. Other animals, like raccoons, have a heightened sense of touch. Their paws have elongated sensor pads, and thousands of touch receptors. With these, raccoons can differentiate textures well enough to tell if a fruit is ripe or not, even in the darkest night.


Raccoons have a strong sense of touch for moving around in the dark.
Their elongated paws have thousands of touch receptors to increase the
sensitivity of this sense. On the dorsal (back) side of the raccoon’s paw,
whiskers (vibrissae) on the ends of their digits heighten the sense of touch.
Raccoons don’t even have to touch something to sense it; they have vibrissae (whiskers) on the ends of their digits, above their claws. Whiskers in general are a potent aid to nocturnal animals, whether located on faces, paws, or bodies (remember the naked mole rat’s whiskers on its torso in Take Off Your Coat And Stay A While).

Even plants can be adapted for nocturnal activity. Moonflowers, night-blooming philodendrons, and other flowers that rely on nocturnal pollinators tend to be white (since their pollinators most likely can’t sense color), and strong smelling. Indeed, the increased temperature of the P. selloum spadix (Is It Hot In Here Or Is It Just My Philodendron) is an adaptation to nocturnality.

So why be nocturnal? Anyone who has tried to negotiate an unfamiliar room in the dark knows that being active in the dark brings certain obstacles that must be overcome. There must be distinct advantages to it or needs for it, or else nature wouldn’t go to the trouble of adapting. Some scientists believe that nocturnality arose from originally diurnal organisms taking advantage of an underused ecological niche. Being active at night can be a form of crypsis (hiding), either to make them better hunters, or to avoid being hunted.

Nocturnality can also reduce the amount of water lost to the environment, and can lower the thermal stress on certain species of animals. For example, many frogs lose water through their skin, so daylight and higher temperatures can dehydrate them quickly.

That doesn’t mean that certain species won’t be exceptions. Moths are all nocturnal, except for the polka-dotted wasp moth, that is. There are four species of wasp moths, all diurnal, but the polka-dot is the prettiest, so we will fall into that old trap and give the pretty one all the attention. Diurnally active, this moth has abandoned many of the nocturnal adaptations of its brethren.


The polka dot moth has color and patterns that might be useful
for mating or for warding off other animals, but they would
be wasted if the animal was nocturnal.
For instance, it is beautifully colorful, a no-no for nocturnal moths. Since color doesn’t show up at night, moths are generally white, tan, or grey. Second, the coloration, especially the bright rump, mimics a wasp (hence the name) and warns of a toxic mouthful if consumed. This defense is called aposematism (apo = away from, and soma = body, basically, keep away from me). Many brightly colored insects will make predators sick, purely a diurnal method of survival, as the warning colors would be of no use at night.

Just as this moth species is diurnal when its close relatives are nocturnal, there is a single genus of primate that has chosen to be nocturnal when all others, including humans, are diurnal. Owl monkeys (8 species) live in Central and South America, and leave their sleeping sites about 15 minutes after sunset each day. They forage for fruits and the odd flower or insect until just before sunrise, then retreat to a hollow tree or within dense foliage to sleep away the day.

Owl monkeys adopted a nocturnal pattern after millions of years being diurnal, so it must have afforded them some advantage or was an answer to some overwhelming stressor. They have adapted by developing larger eyes, with more rods and fewer cones. They still see color, but less so than other monkeys.


The owl monkey is nocturnal, so it needs to have more sensitive vision.
For this reason, it eyes (and eye sockets) are huge! Compare the eye
size and skull morphology in the diurnal capuchian monkey. Form of
the skull follows the functional capacity of the eye.
Owl monkeys are interesting to science for being the source of another exception, as they are the only primates susceptible to the human form of malaria. In The Perils of Plant Monogamy, we used malaria in chimps and humans as an example of divergent evolution; malaria developed into species-specific forms. But the owl monkey is susceptible to both the primate and human species, so they can substitute for humans in malaria research.

Malaria is caused by a parasite, and as such, depends on its host organism for nutrition. The rule is that parasites are active when their host is active (feeding). A good example is the intestinal parasite of the surgeonfish, E. fishelsoni (Of Fish Guts And Cancer).

As I am sure you have committed to memory and made a part of your life, E. fishelsoni grows to an amazing size and replicates its DNA thousands of times before it divides into two or three progeny organisms. It takes tremendous energy for a bacterium to grow 80 fold and produce 85,000 copies of its DNA in one day, so it must occur when nutrients and carbohydrates are plentiful - during the day when the fish is feeding. Although it is a stretch, I guess you could call E. fishelsoni a diurnal parasite.

The malaria parasite, Plasmodium falciparum, has chosen a different path. P. falciparum’s host is man, and man is diurnal (teenagers and third shift workers excepted), but the parasite works to produce many progeny (gametophytes) and have them mature in the nighttime. The reason is simple; malaria has two hosts.


Plasmodium falciparum needs two hosts to complete its life
cycle. One immature form (sporozoite from oocyst) grows
only in the mosquito, while another (gametocyte) forms only
from mature sporozoites in the human red blood cells.
While one stage of the organism grows in the human, another needs to be inside a mosquito in order to complete its life cycle. After finishing its development, it is ready to be injected into another human when the mosquito feeds again. The key is that the mosquito is nocturnal and the gametophyte is short-lived. The gametophyte must be produced and mature just in time to be sucked and deposited into the mosquito gut. P. falciparum has pressured to conform to the activity of one host while it is inside a host with the opposite activity pattern.

It is common that most species within a group will have similar activity patterns, since they are derived from common ancestors and therefore many characteristics are similar, including those that determine fitness for day life or nightlife. But there are exceptions. For instance, most rodents are nocturnal, but we see squirrels all day long - they are diurnal. Also, we mentioned above that most primates are diurnal, but the owl monkeys are nocturnal.

But there are bigger exceptions, organisms that aren’t diurnal or nocturnal. Ants, primates, and cats have species that are all over the place; some are nocturnal, some are diurnal and some are neither. It is the in-betweeners and the neithers that we will talk about next time.


Kreysing, M., Pusch, R., Haverkate, D., Landsberger, M., Engelmann, J., Ruiter, J., Mora-Ferrer, C., Ulbricht, E., Grosche, J., Franze, K., Streif, S., Schumacher, S., Makarov, F., Kacza, J., Guck, J., Wolburg, H., Bowmaker, J., von der Emde, G., Schuster, S., Wagner, H., Reichenbach, A., & Francke, M. (2012). Photonic Crystal Light Collectors in Fish Retina Improve Vision in Turbid Water Science, 336 (6089), 1700-1703 DOI: 10.1126/science.1218072


For more information or classroom activities on activity cycles, night vision or adaptation, see:

diurnal/nocturnal –

night vision –

adaptation –
http://www.nationalgeographic.com/xpeditions/lessons/17/g35/smcreatecreature.html

Do You Have Be Ugly to Hear Well? – Owls and Body Plan Symmetry

Biology concepts – body plan, bilateral symmetry, cephalization, form follows function

Paradox alert – the most complex organisms in nature are the best at reducing complexity. How is that? Nature tends toward symmetry through evolution. Lower organisms do not to have much symmetry, while more complex organisms usually have a symmetric body plan.


Sponges have no symmetry, some lower animals have radial symmetry, while higher animals (including us) have bilateral symmetry. 
Bilateral symmetry results in a front end (anterior) and a back end (posterior).
As far as animals go, sponges have little or no symmetry (yes, sponges are animals). As you move up the ladder of complexity, you first see radial symmetry (starfish, worms, octopuses), then bilateral symmetry (one side of an animal is mirrored by the other side).   

Having a mirror image means no additional planning. It’s like building a second building using the plans from the first - no added cost. By repeating units (metamerization, as in worms and arthropods) or mirroring structures in bilateral symmetry, the animal may become more complex, without having a more complex organizational plan and therefore fewer possible mistakes in development. Paradox averted.

Symmetry leads to distinct front and back ends and movement in a certain direction (see the lobster above). This leads to cephalization (development of a head). Cephalization in turn leads to more bilateral symmetry, including of the head itself. The two halves of the face are close to being mirror images.

Bilateral facial symmetry is thought to be important in determining what people think is pretty. Asymmetries in facial characteristics, even if not noticed consciously, may play a role in determining who we believe to be attractive. For example, men with more facial symmetry have more sexual partners and are more likely to have partners outside their primary relationship….. apparently, symmetry has little to do with decency.


The symmetry of different aspects of the face may be a sign of health.
Facial symmetry might be even more important evolutionarily, as small asymmetries that begin in utero (in the womb) could indicate an inability to resist the harmful effects of environmental or infectious perturbations. This would be a sign of weaker genes and would discourage potential mates. In research on some South American tribes, the more symmetric males tended to have more children survive to adulthood and those children had fewer diseases. It may be that in the search for the healthiest possible mates (therefore the most desirable genes), we use symmetry as a discriminator. Modern medicine has made much of this moot, but instinct is hard to kill.

Now for our exception. One animal has abandoned the move toward symmetry in order to improve its ability to hear. Owls are truly more interested in substance over style; they break the rule of symmetry in order to survive. Nocturnal owls must be able to locate their prey in the dark, and for this they rely on their hearing more than sight, so their auditory sense is truly a survival mechanism.

To hear a mouse, or a ferret, or a cheeseburger from a long distance away is one thing, but owls also need pinpoint from where that sound is coming. Many animals (including humans and owls) are capable of detecting small differences in the time that sounds reach each ear. This is one of the beauties of bilateral symmetry - we have two ears.

If a sound originates from your left, it reaches your left ear before it reaches your right. Your brain senses this time difference and calculates how far left of center the object must be. Humans and owls are equally good at this; we can detect a time difference of less than 10 millionths of a second (0.00001 sec)! In addition, if the sound is closer to your left ear, the sound reaching it will be just a little louder than the sound reaching your right ear. Your brain can sense this difference as well.


There is a large disparity in the vertical position of the ears
in many species of owls. Does it make them unfit or ugly?
However, owls take this a step further, since the third dimension is of more importance in their world. Owls need to know if their prey is above or below them (and by how much) in order to hunt efficiently. With ears at the same height, a sound from below or above reaches both ears at the same time – no help in locating dinner.

On the other hand, if the ears were located at different heights, the sound would reach one ear before the other, and locality information could be obtained. This is what evolution has done for many species of owl. Perhaps not the prettiest solution, but beautiful none-the-less.

Looking at the picture of the owl skull, you can see that the right ear operculum (opening) is placed well above that of the left. This asymmetry does wonders for their sense of hearing, but leaves them with a lopsided head. Lucky for their love life, most owls’ head and facial feathers tend to even out this disparity.
           
The ear asymmetry isn’t the owl’s only body design modification. The combination of the elements listed below allows owls to hear a mouse burrowing under six inches snow up to 100 ft. away, or hear it squeak from a half mile away! It would seem that owls are designed to pick up noise.

Head turn – The asymmetry of the ears and their location on each side of the head allow the owl to localize the sound to a certain degree, but it is increased by the owl turning its head from side to side. The head position where the sound reaches the owls ears at the same time defines when the prey directly in front of its nose. The owl can do this without moving because of its extraordinary ability to turn its head 270 degrees in either direction. This would mean that you could turn your head to the right and end up looking at your left shoulder!
           
Facial disk – In general, the bigger the facial disk on an owl, the more it relies on hearing as opposed to sight to locate prey. The disk is shaped to collect sound and funnel it toward the ears, much the same way that our outer ears collect sound for us, or how a satellite dish collects TV signals. 


The size of the facial disk on an owl gives you an idea how much it relies on hearing to catch its prey. The barn owl on the left and the masked owl in the center rely on hearing most, while the Northern Hawk Owl on the right uses primarily its eyes to hunt.

An owl’s facial muscles change the shape of the facial disk in order to fine tune the sound entering the ears and better locate prey. This is an important hint as to the importance of hearing in some owls; they have muscles to change the shape of their sound collecting face, yet they can’t move their eyes.
           
Spatial auditory mapping - The signals relating to where a sound is coming from are coordinated in the owl’s medulla (an old area of the brain). This part of an owl’s brain is three times as large as a crow’s, and hints that something special may be going on there. The up and down location information generated by the asymmetry of the ear positions is integrated with the left and right information generated from the ear positions on the side of the head and the head turn. The distance is also estimated by the time and intensity differences of sound wave arrival, especially as the head is turned.

For asymmetric owls, the left-right directional cues lie in the interaural (inter = between, and aural = ears) time difference (ITD), while elevation cues are processed via interaural level (loudness) differences (ILD). These two cues are processed in different parts of the brain and then converge to form an aural map. A recent study has compared the size of the auditory nuclei and evolution of these nuclei in asymmetrical- and symmetrical-eared owls. 

The Canadian research team found that all auditory nuclei in asymmetrically-eared owls are larger than the same nuclei in symmetrically-eared owls, even those not involved in ILD, ITD, or converging of signals. They hypothesize that the enlarged nuclei result in increased locating abilities, but also in an extended hearing range in asymmetrically-eared owls. Comparison between phylogenetic trees indicates that increased locating ability preceded an increase in hearing range, and that they both have arisen more than once in different groups.


Since the sound reaches the ears with different qualities, 
the owl can map the sound to a position in his visual field, 
based on time differences (panel b) and intensity differences (panel c). 
This position is mapped directly on to the owl’s visual map.
All this sound information is then translated onto the owl’s visual map in its brain (the location of objects in space as the owl sees it – most mammals have one of these). The owl actually sees the position of the sound, as if the little mouse’s voice was a big red flag waving at the owl. This auditory and visual cooperation must be important for owls, because when they go blind early in life, owls can no longer hunt by sound.

In each example above, the owl has manifested a functional ability, and in each instance this ability has been honed to increase that function. The form may not be elegant, it may not be easy, it may break a rule of biology, but it must be the way it is to perform its function. This is a basic tenet of biology – form follows function. How something looks is more related to its job rather than to the overall esthetics or appeal of the organism. That is why it is so easy for organisms to break rules – function sometimes demands it. Owls are built for hearing, and as a result, they look like owls. Amazing…..and beautiful.

Gutiérrez-Ibáñez, C., Iwaniuk, A., & Wylie, D. (2011). Relative Size of Auditory Pathways in Symmetrically and Asymmetrically Eared Owls Brain, Behavior and Evolution, 78 (4), 286-301 DOI: 10.1159/000330359 

For more information, classroom activities and laboratories, see:

Body plan –

http://faculty.clintoncc.suny.edu/faculty/michael.gregory/files/bio 102/bio 102 lectures/animal diversity/lower invertebrates/sponges.htm


cephalization –

owl hearing –
http://islandwood.org/forkids/owls-at-islandwood/how-can-we-see-owls/owl-ears-1/asymmetrical-ears 
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