Gill, 1872, 1875[which?]
|Star-nosed mole range|
The star-nosed mole (Condylura cristata) is a small mole found in wet low areas of eastern Canada and the northeastern United States, with records extending along the Atlantic coast as far as extreme southeastern Georgia. It is the only member of the tribe Condylurini and the genus Condylura.
The star-nosed mole is easily identified by the 11 pairs of pink fleshy appendages ringing its snout, which is used as a touch organ with more than 25,000 minute sensory receptors, known as Eimer's organs, with which this hamster-sized mole feels its way around. With the help of its Eimer's organs, it may be perfectly poised to detect seismic wave vibrations.
- 1 Description
- 2 Ecology and behavior
- 3 Snout comparison to visual organ
- 4 Snout as related to optimal foraging theory
- 5 References
- 6 External links
The star-nosed mole is covered in thick, blackish-brown, water-repellent fur, and has large, scaled feet and a long, thick tail, which appears to function as a fat storage reserve for the spring breeding season. Adults are 15 to 20 cm (5.9 to 7.9 in) in length, weigh about 55 grams (2 oz), and have 44 teeth. The mole's most distinctive feature is a circle of 22 mobile, pink, fleshy tentacles (called rays) at the end of its snout, from which it derives its name. These are used to identify food by touch, such as worms, insects and crustaceans.
The extremely sensitive star-like structure is covered with minute touch receptors known as Eimer's organs. The nose is about 1 cm in diameter with roughly 25,000 Eimer's organs distributed on 22 appendages. Eimer's organs were first described in the European mole in 1871 by German zoologist Theodor Eimer. Other mole species also possess Eimer's organs, though they are not as specialized or numerous as in the star-nosed mole. Because the star-nosed mole is functionally blind, the snout was long suspected to be used to detect electrical activity in prey animals, though little, if any, empirical support has been found for this hypothesis. The nasal star and dentition of this species appear to be primarily adapted to exploit extremely small prey. A report in the journal Nature gives this animal the title of fastest-eating mammal, taking as short as 120 milliseconds (average: 227 ms) to identify and consume individual food items. Its brain decides in the ultra short time of 8 ms if a prey is edible or not. This speed is at the limit of the speed of neurons.
These moles are also able to smell underwater, accomplished by exhaling air bubbles onto objects or scent trails and then inhaling the bubbles to carry scents back through the nose.
Ecology and behavior
The star-nosed mole lives in wet lowland areas and eats small invertebrates, aquatic insects, worms, mollusks, small amphibians and small fish. It is a good swimmer and can forage along the bottoms of streams and ponds. Like other moles, this animal digs shallow surface tunnels for foraging; often, these tunnels exit underwater. It is active day and night and remains active in winter, when it has been observed tunneling through the snow and swimming in ice-covered streams. Little is known about the social behavior of the species, but it is suspected to be colonial.
The star-nosed mole mates in late winter or early spring, and the female has one litter of typically four or five young in late spring or early summer. However, females are known to have a second litter if their first is unsuccessful. At birth, each offspring is about 5 cm (2 in) long, hairless, and weighs about 1.5 g. Their eyes, ears, and star are all sealed, only opening and becoming useful about 14 days after birth. They become independent after about 30 days, and are fully mature after 10 months. Predators include the red-tailed hawk, great horned owl, various skunks and mustelids, and large fish, as well as domestic cats.
Snout comparison to visual organ
Vanderbilt University neuroscientist Kenneth Catania, who has studied star-nosed moles for 20 years, recently turned his research to the study of star-moles as a route to understanding general principles about how human brains process and represent sensory information. He called star-moles "a gold mine for discoveries about brains and behavior in general—and an unending source of surprises".
Comparing the mole's snout to vision, his research showed that whenever the mole touched potential food it made a sudden movement to position the smallest rays, the twin rays number 11, over the object for repeated rapid touches. He reports: "The similarities with vision were striking. The star movements resembled saccadic eye movements—quick movements of the eyes from one focus point to another—in their speed and time-course. The two 11th rays are over-represented in primary somatosensory cortex relative to their size, just as the small visual fovea in primates—a small region in the center of the eye that yields the sharpest vision—is over-represented in primary visual cortex." He notes that some bats also have an auditory fovea for processing important echolocation frequencies, suggesting that "evolution has repeatedly come to the same solution for constructing a high-acuity sensory system: subdivide the sensory surface into a large, lower-resolution periphery for scanning a wide range of stimuli, and a small, high-resolution area that can be focused on objects of importance".
The star-shaped nose is a unique organ only found on the star-nosed mole. Living as it does, in complete darkness, the star-nosed mole relies heavily on the mechanical information of its remarkable specialized nose to find and identify their invertebrate prey without using sight (since moles have small eyes and a tiny optic nerve). This organ is often recognized by its high sensitivity and reaction speed. In only 8 milliseconds it can decide whether something is edible—in fact, this is one of the fastest responses to a stimulus in the animal kingdom and is the reason why the star-nosed mole was lately recognized in the Guinness Book of World Records as the world’s fastest forager.
Anatomy and Physiology
The star nose is a highly specialized sensory-motor organ shaped by 22 fleshy finger-like appendages, or tendrils, that ring their nostrils and are in constant motion as the mole explores its environment. The star itself is a centimeter across and thus has a diameter slightly smaller than a typical human fingertip. Nevertheless it is much larger than the nose of other mole species, covering 0.92 cm2 (0.14 sq in) per touch compared to 0.11 cm2 (0.02 sq in) covered by the noses of other mole species. This structure is divided into a high resolution central fovea region (the central 11th pair of rays) and less sensitive peripheral areas. In this way the star works as a “tactile eye” where the peripheral rays (1–10 on each side) study the surroundings with erratic saccade-like movements and direct the 11th ray to objects of interest. Just like the primate’s foveating eye.
Regardless of the anatomical position of the star as a distal (protruding or extending) portion of the nose, this is not an olfactory structure nor an extra hand. The appendages do not contain muscles or bones and are not used to manipulate objects or capture prey. They are controlled by tendons by a complex series of muscles that are attached to the skull in order to perform a role that seems to be purely mechanical. For this purpose, the star also contains a remarkably specialized epidermis covered entirely by 25,000 small raised domes or papillae of approximately 30–50 mm (1.2–2.0 in) in diameter. These domes, known as Eimer’s organs, are the only type of receptor organs found in the star of the star-nosed mole which proof that the star-like structure has clearly a mechanical functioning.
Eimer’s organ is a sensory structure also found in nearly all of the approximately 30 species of mole, however neither contain as much as in Condylura. This large amount of specialized receptors makes the star ultrasensitive—about 6 times more sensitive than the human hand, which contains about 17,000 receptors.
Each Eimer’s organ is supplied by a number of primary afferents, thus the star is densely innervated. It is associated with a Merkel cell–neurite complex at the base of the cell column, a lamellated corpuscle in the dermis just below the column and a series of free nerve endings that originate from myelinated fibers in the dermis, run through the central column and end in a ring of terminal swellings just below the outer keratinized skin surface. All 25,000 Eimer’s organs distributed along the surface of the star have this basic structure in all 22 appendages. Nevertheless the fovea region (11th pair of rays), which is shorter in area, has a lower density of these organs—900 Eimer's organs on its surface while some of the lateral rays have over 1500. This may sound contradictory with the fact that this region has higher resolution and an important role in foraging behavior. However, instead of having more sensory organs this fovea region uses a different approach in which a skin surface may be more sensitive to mechanoreceptice input; it has more innervation density. Rays 1 through 9 each had about 4 fibers per Eimer's organ, while rays 10 and 11 had significantly higher innervation densities of 5.6 and 7.1 fibers per organ, respectively, revealing how the sensory periphery is differentially specialized across the star.
The myelinated fibers innervating the 11 rays were photographed and counted from an enlarged photomontage by Catania and colleagues. The total number of myelinated fibers for half of the star ranged from 53,050 to 93–94; hence the total fibers for the entire star vary from roughly 106,000 to 117,000. This means that tactile information from the environment is transmitted to the SNC rapidly. This would be insufficient without an adequate processing system, but in the star-nosed mole, the processing also occurs at a very high speed almost approaching the upper limit at which nervous systems are capable of functioning. The threshold at which the mole can decide whether or not something is edible is of 25 milliseconds: 12 milliseconds to the neurons in the mole’s somatosensory cortex to respond to touch and other 5 milliseconds for motor commands to be conducted back to the star. In comparison, this whole process takes 600 milliseconds in humans.
The importance of the star-like nose in the mole’s lifestyle is evidenced in the somatosensory representation of the nose. Electrophysiological experiments which consist on electrodes placed on the cortex during gently stimulation of the body demonstrated that roughly 52% of the cortex is devoted to the nose. This means that more than half of the brain is dedicated to processing sensory information acquired by this organ, even when the nose itself is only roughly the 10% of the mole’s actual size. Thus, it may be concluded that the nose substitutes for the eyes, with the information from it being processed so as to produce a tactile map of the environment under the mole’s nose. As other mammals, the somatosensory cortex of the star-nosed mole is somatotopically organized such that sensory information from adjacent parts of the nose is processed in adjacent regions of the somatosensory cortex. Therefore the rays are also represented in the brain. The inferior most sensitive pair of rays (11th) had a larger representation on the somatosensory cortex, even when these are the shortest pair of appendages in the nose of the star-nosed mole.
Other important fact of the representation of the star in the cerebral cortex is that each hemisphere had clearly visible set of 11 stripes representing the contralateral star. In some favorable cases, a smaller third set of stripes was also apparent; opposite to other body structures that have a unique representation, with each half of the body represented in the opposite cerebral hemisphere. Thus, opposite to other species, the somatosensory representation of the tactile fovea is not correlated with anatomical parameters but rather is highly correlated with patterns of behavior. Recordings from active neurons in the somatosensory cortex show that most cells (97%) responded to light tactile stimulation with a mean latency of 11.6 milliseconds. Besides a fairly large proportion of these neurons (41%) were inhibited by stimulation of proximate Eimer’s organs outside their excitatory receptive field. Consequently the ability of the star to rapidly determine location and identity of objects is enhanced by small receptive fields and its associated collateral inhibition system that constrains cortical neurons with short latency responses.
Sensitivity to the Mechanical Stimuli
In 1996 Marasco determined that the threshold by which the star-like structure senses the mechanical stimuli depends on which type of the Eimer’s organ was excited. He characterized three main classes of Eimer’s receptors, including one of slow adaptation (Tonic receptor) and two or rapid adaptation (Phasic receptor). The tonic receptor has a response similar to that of a Merkel cell-neurite complex. It has free terminals and is therefore able to detect pressure and texture with a high sensitivity and at a Random Sustained Discharge. The rapid adapting responses include a Paccinian-like response based on a (on-off) response caused by pressure and mechanical vibrations with maximum sensitivity to stimuli at a frequency of 250 Hz. The differences between both rapid responses rely on the fact that one of them only has a response during the compression phase.
Among the receptors described, Marasco identified that there were receptors relatively unresponsive to compressive stimuli but were acutely responsive to any kind of stimulus that brushed or slid across the surface of the nose (Stimuli applied with large displacements and high velocity). In contrast there were other receptors that responded robustly to small magnitude compression of any kind but were not responsive to sweeping stimuli. The receptors that were sensitive to sweeping were maximally activated across a broad range of frequencies, running from 5–150 Hz, at large displacements, ranging from 85–485 μm. Conversely, the receptors that respond to compressive stimuli showed a narrow peak of maximal activity at 250–300 Hz with displacements from 10–28 μm.
Based on the circular organization of the nerve endings and its innervation pattern in Eimer’s organs, Marasco proposed by mapping experiments that nearly all receptors in the star-nosed mole have a preference for a particular direction of applied stimuli. Thus, while one receptor elicits a strong response if compressed in one direction, it may stay “silent” when compressed in another one.
The threshold of velocity at which the receptors responded was examined and it was identified that the minimum velocity at which the cell responds was of 46 mm/s, corresponding to the approximate speed of the nose during foraging behavior.
Transduction of the Mechanical Signal
Taking into account that Eimer’s organ senses mechanical deformation, its mechanism of transduction can be explained in few steps:
- Stimuli cause depolarization of the receptor membrane causing a receptor potential and therefore a current towards the node of Ranvier.
- If the receptor potential in maintained and the generated current is enough to reach the node of Ranvier then the threshold is reached to produce an action potential.
- When the action potential is produced ionic channels are activated and therefore the mechanical impulse is transduced into electrical.
- This signal is carried along the axon until it reaches the SNC where the information is processed.
Although these summarized steps of mechanical transduction give a hint of how the star-nosed mole convert mechanical information into potential actions, the entire mechanism of transduction behind this intricate mechanoreceptor is still unknown and further studies are still required.
Despite the poorly developed eyes, star-nosed moles have an intricate system to detect prey and understand its environment. When the star-nosed moles explore their environment, the star produces brief touches which compress Eimer’s organ against objects or substrate. When foraging, moles search in random patters of touches lasting 20–30 ms. Catania and collages demonstrated that the tactile organ of the star-nosed mole is preferentially innervated by putative light touch fibers. When the outer appendages of the star come into slight contact to a potential food source, the star is quickly shifted so that one or more touches are made with the fovea (the two lower appendages, pair 11th) in order to explore carefully the objects of interest—particularly potential prey. This foraging behavior is exceptionally fast such that the mole may touch between 10 and 15 separate areas of the ground every second, and can locate and consume 8 separate prey items in less than 2 seconds and begin to search for more prey in as little as 120 ms—although the average time is 227 ms.
The sequence described constitutes handling time, in studies made by high-speed video, the mole always foveated to the 11th appendage to exploring a food item. The use of the 11th appendage of the tactile fovea was surprisingly similar to the manner in which eyes explore details of a visual scene.
This star-like nose also enables the mole to smell underwater, something which was previously thought impossible as olfaction required inspired air to convey odorants to the olfactory epithelium. Although the star-like structure is not a chemoreceptor itself it help the star-nosed mole to produce between 8 to 12 small air bubbles per second, each of 0.06 to 0.1 mm, onto objects or scent trails. These bubbles are then drawn back to the nostrils, so that odorant molecules in the air bubbles are wafted over the olfactory receptors.
In 1993 Gould and collages proposed that the star-like proboscis had electroreception and therefore that the mole was able to sense the electric field of their prey prior to its mechanical inspection by its appendages. Through behavioral experiments they demonstrated that moles preferred the simulated electrical field of an earthworm rather than an earthworm in an identical arrangement without an electrical field. Therefore they suggests that the nerve endings in the star’s tentacles are indeed electroreceptors and that the moles move them around constantly to sample the strength of the fields at different targets as they search for prey. Nevertheless this theory fails to be explained physiologically and has not yet accepted in the scientific community; instead the theory proposed by Kenneth Catania in which the nose modality is tactile seems to be more feasible and accepted.
The development of the star like appendages suggests precursors with proto-appendages of an ancestor’s snout which became elevated over successive generations. Although this theory lacks fossil evidence or supporting comparative data, nearly all extant moles have sheets of the of Eimer’s organ making up the epidermis or their snout around their nares. Also, recent studies of Catania and colleagues identified one North American species (Scapanus townsendi) with a set of proto-appendages extending caudally on the snout which have a striking resemblance with the embryonic stages of the star-nosed mole. Scapanus townsendii has only eight subdivisions on its face, fewer than the 22 appendages found on the star-nosed mole. Such change is of common occurrence in evolution and is explained by the advantage of efficiently adding modules to the body plan without need to reinvent the regulatory elements that produce each module. Thus, although the star is unique in its shape and size, it seem feasible that the structure is based on a more ancestral bauplan as it comprises similarities found in a wide range of other moles and also in the molecular structure of other mammals.
The picture that emerges suggests that star-nosed moles although based on a mammalian basis, are an extreme in mammalian evolution, with perhaps the most sensitive mechanosensory system to be found among mammals. The evolutionary process that led to elaboration of this star-like nose is based in 2 theories. One proposes the development of the structure of the star as a consequence of the selective pressure of the wetland habitat in which the star-nosed mole inhabits. Wetlands have a dense population of small insects and exploiting this resource requires a higher resolution sensory surface than of other moles. Thus, a shift to the wetland environment may have provided a selective advantage for a more elaborate sensory structure. Besides, in wild caught moles of many species, the Eimer’s organs show obvious signs of wear and abrasion. The constant and repeated contact with the soil apparently damages the sensory organs, which have a thin keratinized epidermis. Star-nosed moles are the only species that live in the moist, muddy soil of wetlands, therefore a less abrasive soil of this environment also allow this star-shaped structure to evolve.
The second theory of prey profitability explains the foraging speed of the star-nosed mole. Prey profitability or energy gained divided by prey handling time, is an essential variable for estimating the optimal diet. When handling time approaches zero profitability increases dramatically. Due to the small invertebrate animals available in the wetlands, star nosed mole developed handling times as short as 120 ms, therefore the dazzling speed with which it forages counterbalances the low nutritional value of each individual piece of food, and maximizes the time available for finding food. Further, the proximity of the star-shaped nose to the mouth greatly reduces the handling time required before food can be ingested, and is a major factor in how the star-nosed mole can find and eat food so quickly.
Current Applications in Engineering
The study of highly specialized systems often allows better insight into more generalized ones. The star-like structure as a striking feature may reflect a general trend in its “less remarkable” relatives, just us humans. Nowadays little is known about the molecular mechanisms of tactile transduction in mammals and just like the drosophila fly is to genetics or the giant axon squid is to neurobiology; the star-nosed mole may be the model organism for tactile transduction. The proper understanding of this saccade-like system and associated transduction may lead in the future to the development of neural prosthesis. Besides, the outstanding speed and precision at which the mole performs may provide hints into the structural design of intelligent machines that may be entitled to reply artificially the outstanding ability to sense of the star-nosed mole.
According to optimal foraging theory, organisms forage in such a way as to maximize their net energy intake per unit time. In other words, they behave in such a way as to find, capture and consume food containing the most calories while expending the least amount of time possible in doing so. With extremely short handling times for eating very small prey, star-nosed moles can profitably consume foods that are not worth the time or effort of slower animals, and having a food category to themselves is a big advantage. Furthermore, just behind the 11th ray of the star, the star-nosed mole has modified front teeth that form the equivalent of a pair of tweezers. High-speed video shows these specialized teeth are used to pluck tiny prey from the ground. Neuroscientist Kenneth Catania reports, "It is also clear from the behavior that the teeth and the star act as an integrated unit—the 11th rays, located directly in front on the teeth, spread apart as the teeth move forward to grasp small food. Thus, tweezer-like teeth and the exquisitely sensitive star likely evolved together as a means to better find and handle small prey quickly...it appears that the ability to rapidly detect and consume small prey was the major selective advantage that drove the evolution of the star."
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