Economy hits natural hisory museums

Chicago's Field Museum of Natural History will cut it's budget by 15%. The Field is not only among the top museums in the country but a center of research excellence in the biological sciences. In an article in Nature News Field Museum associate curator and ornithologist Shannon Hackett worries that severe cuts could jeopardize the museum's stature as a leading research center. Hackett comments, "Once you lose your academic stature, it is very difficult to regain". A major problem lies in loses in the museum's endowment dropping from $320 million in the spring to $215 million in November. Museums around the country are experiencing similar problems. Especially hard hit are those institutions that receive state funding in those states with falling tax revenues. The University of Pennsylvania's Museum of Archaeology and Anthropology has cut 18 research positions and the Virginia Museum of Natural History announced in October that the state has ordered a 10% budget cut resulting in job losses and a reduction in hours the museum is available to the public.

Clearly it is as important now as ever to support your local natural history museum. Visit your museum regularly, consider a membership and if you have the means, donate, so that these vital centers of scientific research and education can continue to grow and thrive. The long term economic health of the US is critically linked to a scientifically literate society where discovery and innovation spawns economic opportunity. Natural history museums are key players in building the scientific literacy required in any successful modern economy.

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Pachyderm Problems for Zoo Elephants

ResearchBlogging.orgIn the December 12 issue of the journal Science Ros Clubb of the Royal Society for the Protection of Animals and colleagues, including well known elephant researcher Cynthia Moss, report that captive elephants do not live as long as their free-living counterparts, or even as long as working elephants in Burmese timber camps.

Collecting data from over 4,500 elephants from European Zoos, wild populations in Amboseli National Park in Kenya and working elephants in Burmese logging camps, the authors found a significant correlation between captivity and longevity. Females from a well studied population of African Savannah Elephants (Loxodonta africana) in Amboseli National Park in Kenya exhibited a median life span of 56.0 years (these data excluded mortality from humans). African Savannah Elephants in zoos have a median life span of only 16.9 years. As of 2005 when the study ended female African Savannah Elephants in captivity experienced a mortality risk 2.8 times higher than the natural mortality of wild female elephants in Amboseli. Captive-born female African Savannah Elephants die earlier in zoos than in the wild but infant and juvenile mortality was similar between wild and captive elephants.

For Asian Elephants (Elephas maximus) the effect of zoo captivity on mortality was also significant. Captive female Asian Elephants in the study exhibited a median life span of 18.9 years while working Asian Elephants in a Burmese timber operation had a median life span of 41.7 years. While mortality risk in African Savannah Elephants went down over time, suggesting improved captive management, there was no significant reduction in mortality for Asian Elephants. Also, being born in a zoo versus born in the wild had a significant effect on surviorship in Asian Elephants. Ironically, wild-caught Asian Elephants did better in captivity than their captive-born counterparts.

Elephants live in tightly knit social groups of females and juveniles with very long-term associations among individuals. Wild female elephants rarely move between groups, but, zoos regularly transfer individuals among institutions. Female Asian Elephants are moved around among European Zoos approximately once every 7-years. Transfers have an effect on the health of captive elephants. This study found that inter-zoo transfers significantly reduced survivorship in Asian Elephants.

Georgia Mason, a co-author on this study and zoologist at the University of Guelph in Ontario, Canada, discussed the results on the December 12 Science Magazine podcast. According to Mason the situation for American zoo elephants is no better than their European counterparts. 15% of zoo-born elephants in Europe die in their first year while in the USA 40% of zoo-born elephants die before the age of one.

Small group size, frequent inter-zoo transfers, and comparatively tiny enclosures for an animal that has orders magnitude greater home range area in the wild are all likely contributors to the problem of reduced survivorship in zoo elephants. However, solutions to this problem are not straightforward. Large sums of money have been spent in European and US zoos to build larger enclosures for captive elephants but the study by Clubb and colleagues found little evidence that such improvements have resulted in increased survivorship in captive elephants. Some increases in survivorship for African Savannah Elephants have occurred but not nearly enough to bring their surviorship on par with wild counterparts and the study found that despite increased spending and larger enclosures there was no increase in survivorship for Asian Elephants. Mason in the Science Podcast interview pointed out that recent expenditures of approximately 23 million US dollars spent on improving enclosures for the elephants at the Oklahoma City Zoo were greater than the entire annual budget for the Kenya Wildlife Service or the South African National Parks Authority. Perhaps the greatest concern is that captive elephant populations are not self sustaining and can not survive without introduction of individuals taken from the wild.

This study provides a compelling argument for an elevated discussion on not just captive elephants but the welfare of other large-ranging, social mammals as well. Hopefully this study will place a renewed emphasis on future research and novel approaches to captive husbandry of these magnificent mammals.

Clubb, R., Rowcliffe, M., Lee, P., Mar, K. U., Moss, C., Mason, G. J. (2008). Compromised Survivorship in Zoo Elephants Science, 322 (5908) DOI: 10.1126/science.1164298

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Interview with Judge John E. Jones III

The journal PLoS Genetics has published a wonderful interview with Judge John E. Jones III. A conservative federal judge in the US District Court for the Middle District of Pennsylvania, Judge Jones was recommended for his current position by PA senator Rick Santorum and appointed to the bench by President George W. Bush in 2002. Judge Jones ruled for the plaintiffs in Kitzmiller versus Dover Area School District striking down a school board policy exposing students to intelligent design creationism in the public classroom.

In the PLoS Genetics interview Judge Jones is asked about his own personal views on creationism and evolution. After saying that as a judge he can review a case independent of his personal views he adds,

"I am a person of faith. I'm certainly not an atheist or an agnostic and I see some divine force somewhere. That said, having had a pretty good education, a great liberal arts education at Dickinson College, I must say that I never had any substantial doubts about evolution generally. I had forgotten, admittedly, a lot of what I had learned about evolution back in college. Moreover, a lot had happened since the '70s, so my understanding was rudimentary. But I never had a crisis of confidence about evolution or a reason to doubt that it constituted a valid theory and good science."

Brown University biologist Ken Miller was an expert witness in the trail and recently gave a wonderful lecture on evolution and intelligent design as part of the Cincinnati Museum Center's Dury Science Lecture Series. Here's what Judge Jones had to say about Dr. Miller's testimony in the PLoS Genetics interview,

"I will always remember Ken Miller's testimony in the sense that he did A–Z evolution. And then got into intelligent design. And having laid the foundation with the description of evolution, got into why intelligent design doesn't work as science, to the point where it is predominantly a religious concept."

Contrast this with Judge Jones opinions of the expert witnesses for the defense, particularly Lehigh University professor Michael Behe,

"Another remarkable moment on the science side was Michael Behe, who was the lead witness for the defendants, and a very amiable fellow, as was Ken Miller, but unlike Miller, in my view, Professor Behe did not distinguish himself. He did not hold up well on cross-examination."

That a conservative self-proclaimed "person of faith" appointed to the bench by President Bush rules so desisively against the teaching of intelligent design creationism speaks volumes for the validity of the evolution position and the vacuity of intelligent design from a scientific and, in so much as the idea is applied to public education policy, legal stance.

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Yeast goes with the FLO

ResearchBlogging.org“Share and share alike”, so goes the old adage about equally distributing goods, even at one’s own expense. Darwin’s idea of natural selection explained evolutionary change by posing that traits spread in the population due to the benefits these traits confer on their bearers. Individuals with certain characteristics produce more offspring relative to those individuals with other characters and as a result the population as a whole takes on a different appearance. However, Darwin’s idea couldn’t explain those traits that provide benefits to others at the expense of their bearers. Sharing with others at one’s own expense is called altruism and in biology altruism remained a puzzle for over a century after Darwin.

Then come the 1960’s and William Hamilton. Hamilton said that a rare gene underlying some altruistic behavior could spread despite the cost to the bearer. But how? How could a gene that results in fewer offspring for its bearer spread in the population? Lucky for Hamilton he had an understanding of genetic inheritance that was not available to Darwin. If behavior is directed towards those individuals in the population who also harbor the same genes for altruism then the trait will spread in the population through the recipients of altruistic behavior. Rare genes for altruism would more likely spread if there were a readily observable marker of the same altruistic gene in others. Richard Dawkins popularized this idea in the 1970’s calling it the ‘green beard effect’. In a hypothetical example Dawkins imagined that genes for altruism would result in a marker, such as a green beard, along with the altruistic behavior. With this clear marker of the altruistic gene in others helpers could direct their behavior only towards those that shared the gene for altruism.

Amazingly there is good evidence for the ‘green beard effect’ in nature in several organisms, from slime molds and fire ants. Even more astonishing, there are examples where a single gene is responsible for ‘green beard’ altruism. Scott Smukalla, Marina Caldara, and Nathalie Pochet of Harvard University and their colleagues report in the latest issue of the journal Cell that they have found a ‘green beard’ gene in the budding or brewer’s yeast, Saccharomyces cerevisiae.Yeast is not only the critical component in the making alcoholic beverages but it is also a classic model system in the study of the eukaryotic cell. Yeasts are single celled organisms but wild strains of Saccharomyces cerevisiae in times of stress will aggregate into multicellular mats often called biofilms. These aggregates of cells can protect cells from antibiotics, heat and cold stress, ethanol, and other toxins. The coming together of single yeast cells into a multicellular group is called flocculation and the aggregations are known as flocs. Occurring in wild yeast in response to stress, flocculation allows the population to ride out tough times.

Typical of many organisms grown under the resource-rich and stress-free conditions of the laboratory, times are seldom that tough and many years of culture in the lab have lead to the loss of flocculation in laboratory strains. Comparing a wild, flocculent strain called EM93 with a laboratory strain, S288C, incapable of forming flocs, Smukalla and colleagues found that flocculation fell under the control of a single variable gene called FLO1. This was confirmed by activating the expression of FLO1 in normally non-flocculent S288C cells. Expression of FLO1 resulted in flocculation exactly like that observed in wild yeast. FLO1 expression creates cell membrane proteins that allow cells to recognize and adhere to other yeast cells expressing the FLO1 gene.

FLO1 in Saccharomyces cerevisiae acts like the ‘green beard’ gene predicted by Hamilton as it allows yeast cells to detect others also expressing FLO1 and form multicellular aggregates and thus provide group protection against environmental toxins. But, remember altruism by definition involves a cost to the altruist. Where is the cost? When grown under toxin free conditions and ideal temperatures yeast expressing the FLO1 gene suffer a 4-fold reduction in population growth relative to yeast cultures that do not express the FLO1 gene.

A mixed culture of FLO1 expressing and non-FLO1-expressing cells grown under conditions that lead to flocs results in flocs containing primarily FLO1 expressing cells and free cells that do not express the FLO1 gene. FLO1 is therefore a true ‘green beard’ gene as it promotes the altruistic, social trait (flocculation) and at the same time excludes participation of those cells not expressing the social trait. Requiring FLO1 for cell adhesion eliminates the spread of selfish cheaters, yeast cells that forego the cost of expressing FLO1 while times are good but also reap the benefits of flocs when times are tough.

Research on the evolution of social, altruistic traits like flocculation can shed light on one of the most important transitions in the history of life, the evolution of multicellular organisms from single celled organisms. Saccharomyces cerevisiae, social amoebae, slime molds and many social bacteria move between a single celled and a multicellular lifestyle. Like Saccharomyces cerevisiae, multicellular forms in other microorganisms are often in response to stressful environments. Very early in our own evolution the colonization of harsh environments by our single celled ancestors likely promoted the same altruistic behavior seen in many modern microorganisms today.

S SMUKALLA, M CALDARA, N POCHET, A BEAUVAIS, S GUADAGNINI, C YAN, M VINCES, A JANSEN, M PREVOST, J LATGE (2008). FLO1 Is a Variable Green Beard Gene that Drives Biofilm-like Cooperation in Budding Yeast Cell, 135 (4), 726-737 DOI: 10.1016/j.cell.2008.09.037

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Ken Miller speaks at Cincinnati Museum Center

This Thursday, December 4th at 7:30pm at Cincinnati Museum Center's Union Terminal Dr. Kenneth Miller of Brown University will speak in the museum's Dury Science Lecture Series. Dr. Miller is a cell biologist and author of life sciences textbooks and popular books on the conflicts between evolution and intelligent design creationism.

A vocal proponent of evolutionary biology in the public arena Dr. Miller has been featured on the PBS series Evolution and served as an expert witness in the recent Kitzmiller v. Dover Area School Board case in US District Court of Middle Pennsylvania. Dr. Miller's testimony played a critical role in the Judge John Jones' decision which declared that the Dover school board's policy to promote intelligent design (ID) theory in public school science courses was in violation of the US Constitution's Establishment Clause because, while presented as science, "ID cannot uncouple itself from its creationist, and thus religious, antecedents".

While proving to be highly effective at exposing the deep flaws within this latest version of scientific creationism, ID theory, Dr. Miller has provided a counterpoint to creationist/ID accusations that evolution promotes atheism by being very open about his own personal beliefs in the Christian faith. Miller's highly successful popular book Finding Darwin's God was not only an effective rebuttal against ID but also a personal testimonial of a Christian scientist's ability to reconcile faith with the scientific consensus on biological evolution. His latest book, Evolution and the Battle for America's Soul, dismantles ID arguments and emphasizes the explanatory power of evolutionary biology in making sense of life's diversity.

In a dwindling economy with environmental and energy crises looming larger and innumerable challenges for the US both domestic and abroad the ability to innovate is critical in moving America forward. The launch of the Sputnik satellite in 1957 by the Soviet Union was a wake-up call for the US and spawned an increased focus in science education and research in the 1960's. The rising prominence of China and India in science and engineering is today's Sputnik and Americans need to decide if they want to continue to be leaders and producers of science innovation and technology or followers and consumers of technology provided by other nations.

Far from an esoteric issue the debate between ID creationism and evolution cuts to the heart of science education in the US. On the one side is scientific innovation and adoption of evidence based inquiry and the other is an attempt to roll back two centuries of scientific progress and judge scientific evidence on the basis of narrowly-defined, preconceived socio-religious ideology. Attending Dr. Miller's lecture at the museum center will be an excellent introduction to these critical issues in American scientific literacy and demonstrate that, contrary to what creationists would have us believe, scientific progress need not come at the expense of our religious faith. I hope to see you there.

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DNA lab is coming together


The new molecular genetics lab at Cincinnati Museum Center is slowly coming together. Supported by a grant from the National Science Foundation this lab will provide the instruments needed to apply molecular genetic tools in research in ecology, evolutionary biology and molecular systematics. The centerpiece of the lab is an automated capillary electrophoresis machine, also known as an automated DNA sequencer, manufactured by Applied Biosystems. We received a new lab bench to support this valuable instrument and provide much needed workspace. We also had to run a new electrical line to supply 220V, 30A power to the sequencer. Researchers from partner institutions such as Xavier University and Thomas More College will be touring the new lab soon and start bringing in students to gain valuable skills in cutting edge molecular genetic tools. The lab is getting some attention in the community as well and a story by staff reporter James Ritchie appears in the Buisness Courier of Cincinnati.

Yesterday I ran the very first polymerase chain reaction EVER at Cincinnati Museum Center. This technique used to copy a specific region of an organism's genome is the backbone of modern molecular genetics. Our first reactions were a test run of some genetic sexing reactions amplifying a seqment of the sex-linked chromo-helicase-DNA-binding gene in Red-shouldered Hawks. This work is done in collaboration with Cheryl Dykstra to learn about growth and development of Red-shouldered Hawk nestlings in Southern Ohio.

I have high hopes for the lab. Keep checking back here and at cincyevolution for more updates on this new line of research going on in molecular ecology and systematics at Cincinnati Museum Center.

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How to collect a blood sample from a bird

Genetic tools are a major part of modern comparative biology. Museums are the place where much of this sort of research occurs as museums are storehouses of the source material used in comparative genetic studies. Frozen tissue is stored in ultralow freezers or in liquid nitrogen for a variety of organisms comprise genetic resource collections or GRCs. Ideally tissue samples should be associated with a voucher specimen such as skeletal material, a stuffed skin or a pickled specimen. This allows researchers to check the identity of the tissue sample or compare genetic data with morphological data derived from the source specimen. In ornithology there is a growing trend to collect a blood or feather sample from a bird and release the bird back into the wild. A digital photo together with carefully taken morphological measurements can serve as the voucher for the blood sample. While this is not the "gold standard" way to build a bird collection it can augment traditional collecting efforts and increase numbers of samples while minimizing the effect of collecting on avian populations. Below is a video of me collecting a blood sample from a House Sparrow (Passer domesticus) caught just outside the Geier Collections and Research Center at Cincinnati Museum Center. When done properly there is no evidence of adverse effects on the bird, even though surely it is not an experience they enjoy!

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New National Science Foundation funded DNA lab at CMC!

Fantasic news for zoology research and education at the Cincinnati Museum Center. We just received an award from the National Science Foundation to fund the purchase of instruments for a molecular ecology and systematics laboratory in the zoology department here at the Cincinnati Museum Center! The centerpiece of this new laboratory will be an automated capillary electrophoresis machine. This piece of equipment will allow for DNA sequencing and multilocus genotyping to be done completely in house at Cincinnati Museum Center's zoology department. Numerous projects are already planned to be conducted in the new lab covering a diverse array of topics, everything from DNA barcoding of Neotropical land snails to the population genetics of owls to amphibian conservation genetics to characterizing the genetic mating system of songbirds. The new lab will have close partners in the region including the Cincinnati Zoo, Thomas More College, Cincinnati Country Day School and Xavier University and will facilitate research, future funding opportunites and educational experiences in cutting edge life sciences techniques for high school students and educators, undergraduates, graduate students, local college and university faculty and our dedicated volunteer staff.

Now I just need to think of a name for the lab! I was thinking of the CincyMolES Lab (Cincinnati Museum Center Molecular Ecology and Systematics Laboratory). We could have a Star-nosed Mole as our mascot maybe? OK, I don't study mammals but it's a neat critter and I thought the name was catchy! I'm happy to field other suggestions from my readers.

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AOU/COS/SCO 2008 Meeting in Portland: Part II

Last week was the joint conference of the American Ornithologist's Union, the Cooper Ornithological Society and the Society of Canadian Ornithologists in Portland, Oregon. There were many excellent presentations on a variety of topics in ornithology. Below are, in my opinion, some of the highlights from the conference representing the cutting edge of avian research in North America.

- Rosemary Grant of Princeton University presented a plenary lecture summarizing her collaborative work with her husband, Peter Grant, and numerous students and post-docs on the evolutionary biology and ecology of Darwin's Finches in the Galapagos Islands. The Grants' work on selection on bill size in Darwin's Finches is a classic work, arguably the most famous and well supported examples of Darwinian natural selection in the wild. Now the Grants are tackling speciation and presenting their ideas on how Darwin's Finch species arise over time. Whether or not many currently recognized species of Darwin's Finch are indeed true evolutionary species or complex, polymorphic populations was a hot topic of debate behind the scenes in the lobbies of the Hilton and the bars and restaurants on the streets of Portland. It remains to be seen how the Grants' ideas on speciation will stand up to the scrutiny of their ornithological peers.

- Terry Chesser of the National Museum of Natural History along with co-authors from museums around the country such as the University of Kansas and Louisiana State University presented new data on the relationships of one of the largest and most problematic groups of passerine birds, the Neotropical ovenbirds of the family Furnariidae. The woodcreepers were found to be monophyletic (meaning all the birds currently assigned to the woodcreeper group were found to share a single common ancestor). Two main groups were distinguished within the Furnariidae the woodcreepers and the "true" furnariids, or simply the rest of the furnariids.

- Townsend Peterson of the University of Kansas discussed the power of museums in understanding the spread of animal-borne diseases (aka zoonotic diseases). The "bird-flu" or H5N1 virus has the potential for a global pandemic and since 2003 has persisted as a lingering threat to human health, especially in south and south-east Asia. Peterson's group found in surveys of birds collected from south China that H5N1 is not exclusively a disease found in galliform birds (chickens, etc.) and waterfowl but is also found among passerines (i.e. songbirds) and other wild landbirds. Peterson also used bird banding databases to model the potential outbreak of zoonotic, bird-borne diseases, like H5N1, in North America and found that where the outbreak starts can have dramatics effects on the spread and geography of the subsequent spread. All of these findings can only be possible with large museum-led databases of both specimens and large scale sighting and banding records.

- John Wieczorek of the University of California at Berkeley presented an update on the ORNIS database. ORNIS is a distributed database that links together avian data from dozens of museum collections. Today the ORNIS database allows one not only to access holdings on avian specimens but also millions of sighting and banding records, digital photos and sound recordings. Wieczorek also discussed a new online tool, BioGeomancer, to add georeferencing data to existing specimen, photo, recording and sighting records. The power of these online, distributed databases and the georeferencing tools that accompany them was seen in numerous talks during this meeting. With adoption of it's new KE EMu database Cincinnati Museum Center is poised to join these other institutions in making it's collection more accessible and usable by researchers in novel and powerful ways.

- An entire session was devoted to the hot topic of subspecies in ornithology. The traditional way in which scientists name species is the binomial, the familiar genus and species names given to every organism. However, there is variation within species and to incorporate this variation into taxonomy many have adopted a trinomial system of nomenclature consisting of three names for each species, genus, species and subspecies. Kevin Winker of the University of Alaska at Fairbanks gave a good history of the use of trinomial nomclature in ornithology and discussed why divergence in phenotpyes (those characteristics of an organisms most often accessible to direct observation such as size, color, etc.) is not also equal to genetic divergence. This means that often the readily observable, physical traits used to distinguish between different subspecies of birds may not correspond to much genetic variation and thus may be of little evolutionary significance when it comes to distinguishing independent genetic lineages. Susan Haig of the USGS Forest and Rangeland Ecosystem Science Center discussed the problems that differing species concepts and subspecies designations pose for conservation efforts. Haig believes that subspecies designations are critical for conservation and law enforcement purposes even if their biological footing is less than solid. These points were hotly debated in the session and in discussions outside the talks.

- Perhaps my favorite talk of the entire conference was by Matt Carling and Rob Brumfield of the Louisiana State University Museum of Natural History. An idea in evolutionary biology known as Haldane's Rule says that in hybrids the sex that contains two different sex chromosomes (the heterogametic sex) should be less fit than the sex that contains two of the same sex chromosomes (the homogametic sex). A bad version of a gene can be overruled by the action of a good version of the same gene on another chromosome. Therefore heterogametic hybrids will be affected by all deleterious alleles on a sex chromosome. Haldane's Rule has been shown to predict the weaker sex in hybrids in everything from insects to mammals. In humans, and most other mammals, males are the heterogametic sex by virtue of having X and Y sex chromosomes while females are the homogametic sex having two X chromosomes. However, in birds females are the heterogametic sex with W and Z chromosomes and males with two Z chromosomes. Lazuli and Indigo Buntings hybridize in the central US. Carling and Brumfield measured the change in gene frequency across a hybrid zone between Lazuli and Indigo Buntings for both nuclear genes and genes linked to the sex chromosomes. They found much smaller genetic clines for sex-linked loci than for autosomal loci (those loci on chromosomes other than sex chromosomes). This is consistent with sex linked genetic incompatibilities in the hybrids. They also identified one locus in particular that contributed very heavily to this narrow genetic cline in Lazuli and Indigo Buntings. This loci was matched to a similar sequence in the chicken genome which, in chickens, contributes to the failure to lay eggs. Essentially Carling and Brumfield have identified an important gene which contributes to the maintenance of Lazuli and Indigo Buntings as discrete species. An amazing study which I hope to hear more about in the future.

- A fantastic symposium was presented on the last day of the conference in which some of the top curators and collections managers in ornithology presented techniques on preparing and managing material in avian collections. University of Kansas curator Mark Robbins and University of Washington Professor Emeritus Sievert Rohwer presented a live demonstration of their study skin preparation techniques. With some relatively minor differences our volunteers in the ornithology collection at Cincinnati Museum Center are preparing specimens like the pros do it, however, I did pick up some useful tips that will improve our specimens. Kimberly Bostwick of the Cornell University Museum of Vertebrates demonstrated her techniques for preparing skeletal material from bird specimens, again as a live demonstration. This was also very useful as here at Cincinnati Museum Center we plan on a major push towards increasing the collection's avian skeletal holdings. Also, Kevin Winker of the University of Alaska at Fairbanks presented a general overview of the tools of the trade for avian collecting from the intricacies of the permitting process to auxillary barrels for a shotgun! All of the talks in this session provided useful tips for the growing bird collection at Cincinnati Museum Center but they also reaffirmed that the changes implemented in our protocols have put us on the right track towards having a world class ornithology collection.

-Finally Irby Lovette of Cornell University presented some interesting, if not troubling, results on measuring genetic diversity using molecular genetic markers. Researchers for various reasons often want to know the genetic diversity of an individual organism. Genetic diversity in an individual is typically measured in terms of heterozygosity. At any particular location in an individual's genome (at least for sexually reproducing organisms) there will be two copies of a particular gene. If those two copies are the same the individual is homozygous at that locus if they differ then the individual is heterozygous a that locus. Researchers use different types of genetic markers to determine the degree to which an individual is heterozygous across it's entire genome. However, Lovette demonstrated that different markers do not agree with one another on genome-wide heterozygosity. A level of heterozygosity determined using one type of marker may not correspond to the same level determined by another marker. Even multiple loci of the same types of markers often do not agree. This work is consistent with previous findings showing that levels of relatedness and inbreeding were only evident using very large numbers of genetic loci and that molecular genetic markers were inferior to good pedigree data in determining genetic diversity. However, there are many correlations in birds between molecular genetic based genetic diversity and various fitness measures, such as hatching success or growth or behavioral measures. The question then is what do these correlations mean? This will be a fertile field for future research for sure!

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