The Beautiful Eimer's Organ

Maikäfer by H. Baluschek

May is the season of the Maikäfer in Germany, and moles become most active. Hence, in this month we celebrate Gustav Heinrich Theodor Eimer's arguably greatest discovery. Even Frau von Welt discussed it, though in fleeting. Theodor Eimer was born in Zürich, Switzerland, on Feb. 22, 1843, and passed away on May 29, 1898, in Tübingen, Southern Germany. His biography remains scant. After an extended search, I gleaned his birthday from the simple plaque on his grave pictured in this tour of Tübingen's Bergfriedhof. Plaque is a misnomer here. It consists of a rectangular steel plate with black lettering painted on gray rust-proofing. The simplicity does not come close to reflect his discovery.

Theodor Eimer aspired a career in comparative zoology and anatomy. He spent his junior faculty years as prosector at Julius-Maximillian's University, Würzburg, Germany, and moved on to hold a chair in zoology at Eberhard-Karl's University, Tübingen, where he led a productive life as an evolutionary biologist, struggling with Darwin's theory until his untimely death at the age of 55. His greatest scientific contribution, however, may have been as an early neuroethologist. Perhaps, it is this contribution that helped Tübingen's university and its three affiliated Max Planck Institutes develop into a hotbed of neuroscience today.

During his tenure in Würzburg, Theodor Eimer became the first to describe the discrete microscopic organ of touch that densely populates the tip of the nose of the European mole Talpia europaea. The organ is named in his honor. In his original publication (Eimer, 1871), he examined in great detail the structure of the nose, the distribution of the touch organs on the nasal skin, and the relationship of their density with the nose's use for palpation. Eimer sought to establish a connection between structure and function.

Eimer's organ fascinated me, ever since I saw one under the microscope in histological preparations Kenneth Catania showed me. Kenny is studying the North American star-nosed mole Condylura cristata. He counted roughly 25,000 Eimer's organs on these moles' nose appendages called rays. The organ consists of a minute skin papilla between a tenth and a fifth of a millimeter in diameter. At the papilla's core, a beautifully geometric constellation of nerve fibers with free endings is embedded with great symmetry in a conspicuous column of epithelial cells. Eimer (1871) saw two to three single nerve fibers, rising strait in the middle of the column and ending in the fifth layer under the stratum corneum that forms the hard top of the epidermis. The fibers extend short protrusions perpendicularly into each epithelial layer they traverse, where the protrusions end in 'buttons'. They are ringed by a circle of roughly 19 evenly-spaced nerve fibers, known as satellite fibers, whose protrusions point inwards. In addition, Eimer (1871) distinguished a separate set of nerve fibers with free nerve endings. By contrast to the fibers in the papilla's core, these travel obliquely toward the surface at the papilla's perimeter. Both sets are illustrated in his Fig. 2 of his 1871 publication, partially reproduced below:

His legend to this figure reads: Longitudinal section through the frontal surface of the mole's snout treated with the Gold-method. T touch cone, N nerve bundle, E epithelium, C corneum, magnification 400/1.

With improved histological techniques, a second touch receptor type, Merkel cell-neurite complexes, was found in the stratum germinativum at the bottom of the epidermis (darkest part in Eimer's figure), and a third, lamellated corpuscles of Vater and Pacini, was discovered in the stratum papillare of the dermis (white in Eimer's figure) underneath the Merkel cells (Halata, 1975).

Today, we still do not understand precisely how these receptors transduce touch into the electrical signals that the nerve fibers transmit to the brain. But we have learned much about the properties of touch, e.g. frequency and force, to which the receptors respond and how their responsiveness changes with prolonged stimulation. The receptors can be functionally distinguished based on these features. The nerve fibers with free nerve endings and the nerve fibers ending on Merkel cells adapt their responses to touch rapidly, whereas the nerve fibers ending in the lamellated corpuscles are considered slowly adapting.

Marasco and others (2006) were able to attribute different functions to Eimer's two sets of free-ending nerve fibers in the star-nosed mole and the coast mole Scapanus orarius. The authors furnish outstanding micrographs of the organ and its innervation, depicting Eimer's free-ending fibers as well as the Merkel cell-neurite complexes and the Vater-Pacini corpuscles. Using a histochemical marker for a protein known to be involved in the processing of pain, they were able to label the nerve fibers at the perimeter of the papilla, suggesting that they are nociceptive. That is, they respond to pain. By contrast, the fibers in papilla's core did not stain for the protein, suggesting that they are mechanoreceptive. These nerve fibers as well as the Merkel cell-neurite complexes are known to respond to local touches with great sensitivity, whereas the Vater-Pacini corpuscles are highly tuned to the frequencies of dispersed vibrations. Eimer's organ, therefore, forms a receptor complex, integrating pain receptors as well as three fundamentally different types of touch receptors which preferentially respond to either skin indentations or vibrations. The follicles of whiskers, also known as vibrissae or sinus hairs, and the push rods in monotremes (Proske and others, 1998) represent the only other known discrete structures in the skin that combine three mechanoreceptor types.

The Eimer's organs on the nose may be the mole's main tool with which the animal can capture a refined picture of its underworld.  Catania and Kaas (1995) have shown that the nose of the star-nosed mole is mapped in multiple topographic representations on an extraordinarily large swath of cerebral cortex that processes touch. Discrete morphological modules of nerve cells that are clearly discernible in histologically stained sections represent each ray in the same order as they surround the nose. This topographic morphological representation of the sensory periphery is similar to that of the facial whiskers by cytoarchitectonic modules called barrels in the rodent cerebral cortex. I touched on whiskers and barrels in my posts dated Oct. 12, 2009, and Dec. 24,  2008.

To date, two complete cortical maps of the nose with its rays have been found in the brain of the star-nosed mole. There may be more. The nose's disproportionate representation in cerebral cortex is suggestive of a fovea for nose touch in the mole's somatic sensory system (Catania, 1995).

Mole - overviewBBC Natural History Unit

Already Professor Eimer recognized the importance of the mole's nose to the behavior of the species. He begins the introduction to his 1871 publication with the statement: "The mole's snout must be the seat of an extraordinarily well developed sense of touch, because it replaces almost entirely the animal's sense of face, constituting its only guide on its paths underground." He estimated that the nose of the European mole was covered with more than 5,000 Eimer's organs which were invested with 105,000 nerve fibers. He took the abundance of sensory innervation to affirm his contention that the nose's touch must represent the moles dominant facial sense. Professor Eimer ends his report with the assertion that his interpretation is entirely consistent with the common knowledge of his time. His last sentence reads: “This enormous richness of innervation easily explains the well-known fact that already a light blow to the snout kills the mole instantly.”

Roughly 130 years after Professor Eimer's discovery, Catania and colleagues accomplished to record striking behavioral evidence in favor of his conclusion, using a high-speed camera that the eminent professor could hardly have imagined  (Catania and Remple, 2004).  Professor Eimer would have been fascinated, watching this footage. Moles with the help of their Eimer's organs may be perfectly poised to detect seismic vibrations.

References

• Catania KC (1995) Magnified cortex in star-nosed moles. Nature 375: 453-454.
• Catania KC, Remple FE (2004) Tactile foveation in the star-nosed mole. Brain Behav Evol 63: 1-12.
  
• Catania KC, Kaas JH (1995) Organization of the somatosensory cortex of the star-nosed mole. J Comp Neurol 351: 549-567.
• Eimer Th (1873) Die Schnauze des Maulwurfs als Tastwerkzeug. Arch Micr Anat 7: 181-201.
• Halata Z (1975) The mechanoreceptors of the mammalian skin ultrastructure and morphological classification. Adv Anat Embryol Cell Biol 50: 3-77.
• Marasco PD, Tsuruda PR, Bautista DM, Julius D, Catania KC (2006) Neuroanatomical evidence for segregation of nerve fibers conveying light touch and pain sensation in Eimer's organ of the mole. Proc Natl Acad Sci USA 103: 9339-9344.
• Proske U, Gregory JE, Iggo A (1998) Sensory receptors in monotremes. Philos Trans R Soc Lond B Biol Sci 353: 1187–1198.

Related Posts

• Force & Resilience
• Neuroanatomy of An Earthquake
• The Quest for the Infrasound Fovea
• The Power of Instruction & The Brain

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Addendum
Impressively shown by the two Reuters news clips below, high-speed video cameras have developed into invaluable tools in the study of animal behavior. Their application range from the examination of bat flight

to feline drinking, providing new insights propelling engineering design (11/12/10).

The Quest for the Infrasound Acoustic Fovea

In my post dated Sep. 30, 2009, I briefly discussed the discovery of echolocation as a means for bats to navigate their environment in total darkness and identify insects to feast on. I noted that in echolocating bats a large part of the auditory system is devoted to the processing of ultrasound, that is frequencies higher than the human audible (>20 kHz).  A disproportionate number of auditory nerve cells are tuned to analyze the frequencies of the echos of the echolocation calls bats emit. I worked on research with echolocating bats in Gerhard Neuweiler's laboratory at the Institute of Zoology of the Johann Wolfgang Goethe University, Frankfurt am Main. I helped show that exposing a bat to its echolocation frequency activated prominent regions in its auditory midbrain structure known as inferior colliculus (Melzer, 1985). The proclivity in the bat auditory system for a narrow band of ultrasound used for echolocation is known as high frequency filter or acoustic fovea (Neuweiler and others, 1980).

As informative contrast to the bats' fovea for extremely high pitch, a mammal was sought with a fovea for extremely low pitch. It was hoped that a similar association between behaviorally relevant sound and its representation in the brain might exist in species whose hearing is particularly sensitive to infrasound, that is at frequencies beneath the human audible (<20 Hz). Elephants (Herbst and others, 2012) and humpbacked whales are known to produce and hear infrasound, but were considered difficult to study.

By contrast, burrowing rodents were thought to constitute suitable candidates, because they spend much of the day underground and were observed to be able to use seismic vibrations to identify and locate conspecifics and predators. Particularly, gerbils were of interest. They are essentially crepuscular, that is most active above ground at dawn and dusk. They live in semi-arid deserts where compacted gravel and sand carry low frequency sound long distances. Gerbils use hind foot drumming as means of communication. The drumming exhibits species-specific differences and has been observed to alert companions to one's own presence in as much as to approaching predators (Randall, 1997). Differences in the signature of the sound may permit the animals to distinguish between various types of predator, e.g. snakes or birds. Moreover, the sounds may convey to predators that they have been discovered (Randall, 2001).

Christian Winter recruited Jürgen Möller as a junior faculty member to investigate whether gerbils may represent the small mammals with an acoustic fovea for infrasound. He had extensive experience in acoustics, micro-electrode recordings of electrical nerve cell discharges, and animal behavior.

courtesy J. Möller

In addition to the well-known Mongolian gerbil (Meriones unguiculatus), Jürgen succeeded in bringing a number of gerbil species from Israel to the laboratory, testing their hearing for low frequency sensitivity with audiograms. Field studies on the animals' behavior were conducted in Israel.

The photograph depicts a sand rat (Psammomys obesus) in the Negev desert at dusk. Sand rats were the largest gerbils in Jürgen's collection.  Power density spectra of their drumming's acoustic frequency components were recorded to examine whether the drumming produced infrasound. Broad spectrum audiograms were constructed from recordings of small voltage changes in the inner ear (cochlear microphonics) and from nerve cell activity in the auditory midbrain (inferior colliculus) to investigate whether the animals could hear infrasound.

The results were presented at a joint symposium of Hebrew University of Jerusalem, Université de Lyon, and Johann Wolfgang von Goethe University entitled "Neurobiology and Strategies of Adaptation". The symposium was convened in Frankfurt am Main in 1981. The two figures below show representative results of the frequencies of the sound produced by drumming (A) and of the sound processed by the auditory system (B). The green band covers the frequency range between 0.5 and 1.0 kHz in which the audiograms (B) show peculiar low frequency sensitivity.

(A) Power Density Spectrum

The frequency of the drumming's acoustic components [kHz] (y-axis) is plotted versus time [ms] (x-axis; courtesy J. Möller).

(B) Audiograms

The sound pressure level [dB SPL] of the most sensitive response (y-axis) is plotted versus sound frequency [kHz] (x-axis). Cochlear microphonics: CM; nerve cell responses in the inferior colliculus: IC (courtesy J. Möller).

Analysis of the power spectra revealed that the animals' drumming contained low frequencies of significant power, infrequently dipping into infrasound. Yet, the audiograms of the animals provided no evidence of an acoustic fovea for such low frequencies. Regardless, the findings clearly suggested that the drumming produced sound to which that the animals' hearing was sensitive.

My project's aim was to visualize pitch-related nerve cell activation in the brain with a functional imaging method (Melzer, 1984). M. Müller (now private docent at the J.W. Goethe University) helped me substantially in this endeavor. The project would not have been possible without the use of the whole-body cryotome in H.-M. Kellner's division at the Hoechst AG's Radiochemical Laboratory.

Animals were exposed to sound of low, medium and high pitch. Pitch is represented tonotopically on the transverse plane through the inferior colliculus. That is, nerve cells in the upward (dorsal) aspect of the structure are particularly sensitive to low frequency sound and become progressively more sensitive to higher frequencies with increasing depth. Functional imaging would reveal the isofrequency domains, permitting us to assess the prominence of low frequency processing in the auditory system.

The figure below shows frequency-related nerve cell activation in transverse slices through the inferior colliculus. Nerve cell activation is coded in pseudo colors (blue: low; red: high). The narrow blue/purple lines near the top are the animals' scalp, that is dorsal is up. The brainstem is at the bottom, that is ventral is down. The animals' left side is on the right.

Neurofunctional Images

The slices were obtained from sand rats exposed to tone pips of (clockwise from top, left) 0.8, 2.5 and 17.0 kHz, respectively. Stimulus-unrelated nerve cell activity was obtained from unexposed animals (bottom, left). The inferior colliculus is the distinctly sound-activated, double-lobed structure at the center of the images. Unexposed animals revealed slightly elevated nerve cell activity in the top aspect of the structure on both sides (bottom, left). Low frequency stimulation produced wide-spread activation, peaking in a band at the top of the structure (top, left). This activation partially overlapped with the observed stimulus-unrelated activity (bottom, left). By contrast, the middle frequency activated a narrow band at mid-depth of the structure on both sides (top, right), whereas activity at the top was almost completely suppressed. High frequency stimulation resulted in similarly narrow bands of activation on both sides. Only in this case, the bands were located at the bottom of the inferior colliculus (bottom, right). In addition, prominent activity was distinct at the top of the structure.

The progression of bands of elevated nerve cell activation from top to bottom with increasing frequency of stimulation is consistent with the notion of a tonotopic map where frequencies are represented in logarithmic progression. Intriguingly, the low part of the sound spectrum was slightly disproportionally represented. However, such disproportionality is not uncommon and has been observed in a number of species (N. Suga, personal communication).

The narrow bands of activation observed with the middle stimulus frequency conformed most distinctly with a representation of frequencies in discrete isofrequency layers. According to the cochlear microphonics, the animals' ear is most sensitive at this frequency. By contrast, the wide-spread activation at the low frequency and the additional foci at the top of the inferior colliculus at the low and the high frequency did not precisely adhere to the principle of discrete frequency representation. We did not know how to interpret these results.

In hindsight, we were perhaps too narrowly focused on auditory responses. Parts of the inferior colliculus are known to receive somatic sensory input. Sand rats have long mystacial whiskers that are in frequent contact with the soil. Cytoarchitectonic structures known as barrels represent the mystacial whiskers topographically in a large swath of the gerbil's cerebral somatic sensory cortex (Rice and others, 1985). Distinct septa separate the barrels. Nerve cells in the septa are known to be particularly attuned to the processing of stimulus frequency (Melzer and others, 2006). Recent observations show that whiskers may resonate (Moore, 2004) to low frequency vibrations, touch receptors in the whisker follicles transduce frequencies up to 1.0 kHz and possibly greater (Gottschaldt and Vahle-Hinz, 1981), and nerve cells in the somatic sensory pathway may process this information (Kleinfeld and others, 2006). Perhaps, gerbils do possess an infrasound fovea after all! Only the processing of this sound may not be strictly auditory.

Addenda

• Watching gerbils around the time of an earthquake may prove insightful. A tiny whiskered burrowing rodent, the Northern Pocket Gopher Thomomys talpoides, survived the catastrophic eruption of Mt. St. Helens in 1980 (11/06/09).
• Caldwell and others (2010) produced an impressive demonstration of the importance of vibrations in vertebrate sensation and behavior. The research feature on National Public Radio's Talk of the Nation (Science Friday) yesterday with the title "Rumble in The Jungle" shows red-eyed treefrogs (Agalychnis callidryas) from the rain forests of central America communicating with vibrations mediated by the twig they sit on. Listen to the podcast of Ira Flatow's conversation with Flora Lichtman entitled "Red-Eyed Treefrogs Rumble in the Jungle". The authors of the study, affiliated with Boston University, contributed excellent footage of the frogs' interactions (05/22/10).

• On Apr. 8, 2011, National Public Radio's Talk of the Nation/Science Friday broadcast an insightful segment with the title "Seeing The World Through Whiskers" on the research of Mitra Hartman and her colleagues (Towal and others, 2011) examining whisking in rats with high-speed video (04/11/2011):

• The magnitude 5.8 earthquake on the East Coast on Aug. 23, 2011, centered near Mineral, Virginia, was felt roughly 120 miles away by animals of the National Zoo in Washington D.C., in some instances tens of minutes before humans did. Listen to Ira Flatow's interview with Brandie Smith, Senior Curator of the National Zoological Park, Smithsonian Institution, Washington D.C., entitled "Did you feel it?" on National Public Radio's Talk of the Nation/Science Friday broadcast today (08/26/2011).
• Listen to Dr. Tecumseh Fitch explaining to Audie Cornish of National Public Radio's All Things Considered in his interview with the title "Study: Humans, Elephants User Similar Vocalizations" broadcast today how elephants produce and may use infrasound (08/09/2012).

References

• Caldwell MS, Johnston GR, McDaniel JG, Warkentin KM (2010) Vibrational signaling in the agonistic interactions of red-eyed treefrogs. Current Biol: doi:10.1016/j.cub.2010.03.069
• Gottschaldt KM, Vahle-Hinz (1981) Merkel cell receptors: structure and transducer function. Science 214:183-186.
• Herbst CT, Stoeger AS, Frey R, Lohscheller J, Titze IR, Gumpenberger M, Fitch WT (2012) How low can you go? Physical production mechanism of elephant infrasonic vocalizations. Science 337: 595-599.
• Kleinfeld D, Ahissar E, Diamond ME (2006) Active sensation: insights from the rodent vibrissa sensorimotor system. Curr Opin Neurobiol 16:435-444.
• Melzer P (1985) A deoxyglucose study on auditory responses in the bat Rhinolophus rouxi. Brain Res Bull 15:677-681.
• Melzer P (1984) The central auditory pathway of the gerbil Psammomys obesus: a deoxyglucose study. Hear Res 15:187-195.
• Melzer P, Sachdev RN, Jenkinson N, Ebner FF (2006) Stimulus frequency processing in awake rat barrel cortex. J Neurosci 26:12198-12205.
• Moore CI (2004) Frequency-dependent processing in the vibrissa sensory system. J Neurophysiol 91:2390-2399.
• Neuweiler G, Bruns V, Schuller G (1980) Ears adapted for the detection of motion, or how echolocating bats have exploited the capacities of the mammalian auditory system. J Acoust Soc Am 68:741-753.
• Randall JA(2001) Evolution and function of drumming as communication in mammals. Am Zoologist 41:1143-1156.
• Randall JA (1997) Species-specific footdrumming in kangaroo rats: Dipodomys ingens, D. deserti, D. spectabilis. Anim Behav 54:1167-1175.
• Rice FL, Gomez C, Barstow C, Burnet A, Sands P (1985) A comparative analysis of the development of the primary somatosensory cortex: interspecies similarities during barrel and laminar development. J Comp Neurol 236:477-495.
• Towal RB, Quist BW, Gopal V, Solomon JH, Hartmann MJZ (2011) The Morphology of the Rat Vibrissal Array: A Model for Quantifying Spatiotemporal Patterns of Whisker-Object Contact. PLoS Comput Biol 7: e1001120. doi:10.1371/journal.pcbi.1001120.

Footnote

• The text of this post is available for download in pdf-format from the scribd store.

Related Posts

• Echolocation, Science & Power
• The Elusive Sense of Magnetism
• Konrad Lorenz, Imprinting and Functional Brain Imaging
• The Power of Instruction & The Brain
• The Brain: A Giant Self-Medicating Gland
• Force & Resilience

I attached this amateur video of a pet gerbil drumming in the cage. The clip provides an idea of drumming speed and periodicity. Keep in mind that a gerbil in a burrow will sound quite different.

Parent Scientists, Children & Informed Consent

On Jan. 17, 2009, Pam Belluck published an article online in the The New York Times with the title "Test Subjects Who Call the Scientist Mom or Dad", in which she describes scientists using their own children in their research. The methods employed were observation of behavior and non-invasive diagnostic techniques, e.g. magnetic resonance imaging (MRI) of brain development and electroencephalography (EEG). In MRI, a scanner maps tiny realignments to high radio-frequency pulses in the spin axes of atomic nuclei inside the body that have been lined up in a strong magnetic field. The measurements are used to reconstruct our body's interior in image slices. With EEG, tiny electrical signals from the nerve cells in the brain are recorded with wire electrodes attached to the surface of the scalp. I conducted such studies as principal investigator. The research is summarized in a chapter of a book entitled "Blindness and Brain Plasticity in Navigation and Object Perception." Some essential findings are described in the posts dated Dec. 31, 2008, and Dec. 9, 2007, on this blog. Both techniques do not pose any known health risks to the participants. I submitted the following comment with the article (comment #9): 

Asking someone to participate in a scientific research study resembles in many ways ancient rites of hospitality. If we are invited to stay with friends, we take it for granted that they shall take good care of us and protect us from harm within their limits.

People who wish to participate in a research study that does not benefit them directly, but furthers scientific knowledge at large, expect to be well informed about the risks that the study may pose. They certainly will expect that the principal investigator is not going to put them knowingly in harms way and that, in case of an accident, everything possible will be done for them within reasonable limits.

Commonly, MRI studies do not pose any risk greater than everyday life for healthy people who do not wear magnetic parts or electronic devices in their bodies. However, the procedure may be quite intimidating, particularly when children are involved. Therefore, it certainly is reassuring for the participants to know that they and/or their children are asked to undergo a procedure that the principal investigator and her/his children have undergone before. Of course, the participants have the right to stop the procedure at any time without any negative consequences and, implicitly, this right extends to the children of principal investigators.

The same rationale applies to EEG and, in principle, to behavior studies. However, while the MRI scanner and the EEG recorder deliver objective results, it is questionable whether parents can be impartial observers of their children's behavior.

Important to all studies with very young children, the question remains to be answered whether the mind of a five-year old has developed sufficiently to understand risk and give informed consent.

Continuity of the Mind

Today the National Convention of the Democratic Party nominated Senator Barack Obama as candidate for the election to the highest office in the United States. His campaign for candidacy took about two years. Though conditions and means have changed decisively, the challenges that have to be met by the hopefuls for this type of position have remained remarkably similar over the past millenium.

Roughly 800 years ago an ambitious young man set out from his home in the pursuit of the highest office of his nation. It was the beginning of a long campaign trail that took him thousands of miles across the land he aspired to govern, learning about differences in culture of a diverse people and gaining insights into their daily problems. The journey was cumbersome and needed elaborate organization. Several hundred aids accompanied him. They had to set up camp, see to his security, and prepare the next moves. His message had to go out. It had to resonate. He had to convey that he understood the people's needs and that, once elected, he was going to help them. The locals had to be entertained. Vast sums of money were spent. Promises were made. Though his journey was educational, the main purpose of the endeavor was to rally support for his cause. He had a powerful and influential opponent and needed the overwhelming endorsement from people of all walks of life in order to win.

He had the professional credentials. He was formally educated to stand up to the challenges of government. He was well prepared in public administration and law. He was fluent in the legal language of his time and proficient in several other languages. However, he needed to learn the language instrumental in his struggle for national leadership: German. The majority of the people in the nation spoke German. The members of the Electoral Council considered themselves German. They met in a German city to elect the new head of state and government, and they wanted a popular leader.

This was a great disadvantage for our candidate. He had grown up in Southern Italy. He was considered a minority, an outsider. His mother was Norman. Though his father was German, our candidate conversed poorly in the high language. A superb command of German was imperative. Therefore, he campaigned much in Germany and made a great effort to learn the language. Eventually, he mastered it so well that he could write poetry. His was a brilliant mind. He dazzled those who met him. He was gregarious, smart and engaging. His sophistication impressed. He had a great gift of endearment. He attained wide popularity, becoming known as the Kid from Apulia after his birthplace in Italy where he would continue to live for many years. Apulia is quite deforested and arid today. In the our candidate's time, the landscape could have easily resembled Turner's vision.

His popularity would eventually give our candidate the edge in the election. His opponent struck a rather dull pose, though he could impress with a fabled pedigree. He was the member of a Saxon dynasty, known as the Wolves. They counted several figures of worldwide acclaim in their ranks. Alas, the Saxon did not show much political skill, reveled in military prowess and was given to adventure. On election day, the Kid from Apulia carried the vote. Though the Council of Electors had only seven members, the effort on our candidate's part was tremendous. Three electors were clergy. Our candidate had a complicated relationship with the Church. He and the Holy See held diametrically opposed views on the separation of church and state and the prerogatives of the two branches. He was to assume a difficult position.

Although he became nominally head of the nation, his governance was riddled with problems. Revenue was hard to collect. Resources beyond his own were never certain. Any decision of national importance needed the support of powerful locals who pursued their own special interests. He spent much time on the road to negotiate support for his ideas. Alliances were ever shifting, and the Holy See never seized to challenge his claims to power. The struggle would consume much of his energy and almost cost him his life. But, ultimately the Kid from Apulia is not remembered for his political achievements, but for his scholarly writings, the poetry he left behind, and his philanthropy.

He founded one of the first universities in continental Europe, the University of Naples. He enjoyed falconry. Some of his accounts on falcon behavior and training are preserved. His modern-minded examination of the subject strikes today's reader in awe. This man was determined, yet flexible. He had excellent observation skills and a brilliant analytical mind. He kept his mind open, willing to learn and embrace the unanticipated and unknown. He understood that even the most powerful can ill afford ignorance. Hopefully, the next President of the United States will be blessed with such abilities.

Who was this man? His name was Frederic. He was crowned Frederic II, Emperor of the Holy Roman Empire of German Nations in A.D. 1220. We do not know much about his looks. Some believe that the horseman statue in the Cathedral of Bamberg is sculpted in his likeness. We are left with a bronze of his beloved hunting companions and his writings about them. The bronze is on display at the Cloisters in New York City. Pages of his treatise on falcon behavior can be seen in the Vatican Library.