Joseph F. Golob MD

• Resident in Surgery, Case Western Reserve University School of Medicine,
• Department of Surgery, MetroHealth Medical Center, Cleveland, Ohio

Often the repercussions of extinctions are not known until the species is gone arthritis medication at walmart buy celebrex 100 mg overnight delivery, and the species is generally irreplaceable in the system arthritis from back injury cheap 100mg celebrex. This may be because the habitat has been altered to the point that the biological system no longer functions properly rheumatoid arthritis early onset discount celebrex 200 mg visa. Another reason for protecting species of concern is for their value as unique genetic resources rheumatoid arthritis gut bacteria buy celebrex 100mg without a prescription. Every species may provide significant information for future use in genetic research and medical practices arthritis nos definition buy discount celebrex 100 mg on-line. Beyond these practical considerations arthritis pain wikipedia generic celebrex 200mg mastercard, perhaps the most compelling reasons for stewardship are the aesthetic and ethical considerations; there is beauty and recreational value inherent in healthy, species-rich ecosystems. The protection of rare, threatened, and endangered species depends on several factors, including increasing scientific knowledge and concerted efforts from government agencies, conservation organizations, educational institutions, private organizations, and individuals. The following section outlines general recommendations to begin to protect the species outlined in this report. Using the Natural Heritage Inventory in the Planning Process One of the main roles of this document is to integrate ecological and conservation information into the planning process. Considering this information early in the planning process allows costly conflicts with rare, threatened and endangered species to be avoided, and protects these resources for future generations. Land use planning establishes guidelines for the kinds of land uses that are suitable in an area and provides a basis for guiding public and private development to benefit communities, the local economy, and the environment. Zoning and subdivision ordinances then set out rules that implement the land use plan. Planning, zoning, and subdivision ordinances are not only valuable tools for urban and suburban areas where development pressures have already affected the use of open space and the integrity of the natural environment, but are also valuable for rural areas where current losses are less pronounced. These areas Beaver County Natural Heritage Inventory Update 2014 ­ Conclusion and General Recommendations / 33 can apply planning to avoid the haphazard losses of valuable regional resources, while still achieving desirable levels of development. Planning for the land use decisions of today and those of the future is an important task and this Natural Heritage Inventory can serve as a useful tool. Pennsylvania Natural Heritage Program staff and expertise are available for additional technical assistance and planning for the conservation of these sites. General Recommendations the following are general recommendations for the protection of the Natural Heritage Areas within the region. Approaches to protecting a Natural Heritage Area are wide ranging, and factors such as land ownership, time constraints, and tools and resource availability should be considered when prioritizing protection of these sites. Opportunities may arise that do not conform to a plan, and the decision on how to manage or protect a natural heritage area may be made on a site by site basis. The following are approaches and recommendations for natural heritage area conservation. Consider conservation initiatives for natural heritage areas on private land ­ Conservation easements protect land while leaving it in private ownership. It can be tailored to the needs of both the landowner and the conservation organization, and it will not be extinguished with new ownership. Tax incentives may apply to conservation easements donated for conservation purposes. Lease and management agreements also allow the landowner to retain ownership and temporarily ensure protection of land. A lease to a land trust or government agency can protect land temporarily and ensure that its conservation values will be maintained. This can be a first step to help a landowner decide if they want to pursue more permanent protection methods. Management agreements require landowners and land trusts to work together to develop a plan for managing resources (such as plant or animal habitat, watersheds, forested areas, or agricultural lands) with the land trust offering technical expertise. Land acquisition by a conservation organization can be at fair market value or as a bargain sale where a purchase price is set below fair market value with tax benefits that reduce or eliminate the disparity. One strategy is to identify areas that may be excellent locations for new county or township parks. Sites that can serve more than one purpose such as wildlife habitat, flood and sediment control, water supply, recreation, and environmental education are ideal. Private lands adjacent to public lands should be examined for acquisition when a natural heritage area is present on either property, and there is a need of additional land to complete protection of the associated natural features. The donation of land entitles the donor to a charitable deduction for the full market value, and it releases the donor from the responsibility of managing the land. If the land is donated because of its conservation value, the land will be permanently protected. Local zoning ordinances are one of the best-known regulatory tools available to municipalities. Examples of zoning ordinances a municipality can adopt include: overlay districts where the boundary is tied to a specific resource or interest such as riverfront protection and floodplains, and zoning to protect stream corridors and other drainage areas using buffer zones. Often it is Beaver County Natural Heritage Inventory Update 2014 ­ Conclusion and General Recommendations / 34 overlooked that zoning can prevent municipal or county-wide development activities which are undesirable to the majority of the residents, and allow for planning that can meet the goals of the county residents. Prepare management plans that address species of concern and natural communities ­ Many of the natural heritage areas that are already protected are in need of additional management plans to ensure the continued existence of the associated natural elements. Site-specific recommendations should be added to existing management plans or new plans should be prepared. Recommendations may include: removal of invasive plant species; leaving the area alone to mature and recover from previous disturbance; creating natural areas within existing parks; limiting land-use practices such as mineral extraction, residential or industrial development, and agriculture; or implementing sustainable forestry practices. For example, some species simply require continued availability of a natural community while others may need specific management practices such as canopy thinning, mowing, or burning to maintain their habitat requirements. Existing parks and conservation lands provide important habitat for plants and animals at both the county level and on a regional scale. For example, these lands may serve as nesting or wintering areas for birds or as stopover areas during migration. Management plans for these areas should emphasize a reduction in activities that fragment habitat. Adjoining landowners should be educated about the importance of their land as it relates to habitat value, especially for species of concern, and agreements should be worked out to minimize activities that may threaten native flora and fauna. Protect bodies of water ­ Protection of reservoirs, wetlands, rivers, and creeks is vital for ensuring the health of human communities and natural ecosystems. Waterways that include natural heritage areas, identified in the Results section of this report, are important, sensitive areas that should be protected. For example, conserving natural areas around watersheds that supply municipal water provides an additional protective buffer around the water supply, maintains habitat for wildlife, and may also provide (low impact) recreation opportunities. Many rare species, unique natural communities, and significant habitats occur in wetlands and water bodies, which are directly dependent on natural hydrological patterns and water quality for their continued existence. Ecosystem processes also provide clean water supplies for human communities and do so at significant cost savings in comparison to water treatment facilities. Therefore, protection of high quality watersheds is the primary way to ensure the viability of natural habitats and water quality. Scrutinize development proposals for their impact on entire watersheds, not just the immediate project area. Cooperative efforts in land use planning among municipal, county, state, and federal agencies, developers, and residents can lessen the impact of development on watersheds. Provide for natural buffers in and around natural heritage areas ­ Development plans should provide for natural buffers between disturbances and critical zones of natural heritage areas. Proposed activities within the Core Habitat of a Natural Heritage Area should be closely scrutinized for potential immediate impacts to the habitat of elements of concern. Proposed activities within the Supporting Landscape of a Natural Heritage Area should be evaluated for potential long-term impacts to habitats of elements of concern, such as water quality, or quantity degradation, or habitat fragmentation. Disturbances may include construction of new roads and utility corridors, nonsustainable timber harvesting, and fragmentation of large pieces of land. Storm runoff from these activities results in the transport of nutrients and sediments into aquatic ecosystems (Trombulak and Frissell, 2000). Vegetated buffers (preferably of Pennsylvania native plant species) help reduce erosion and sedimentation while shading and cooling the water. Preserving water quality in rivers and streams is important to fish as some species, such as brook trout and some darters, are highly sensitive to poor water quality. Sensitive fish are readily lost from streams when water quality starts to decline. Creating or maintaining a vegetated buffer benefits aquatic animal life, provides habitat for other wildlife species, and creates a diversity of habitats along the creek or stream. Reduce fragmentation of the landscape surrounding natural heritage areas ­ Encourage development in sites that have already seen past disturbances (especially mined and heavily timbered areas). Care should be taken to ensure that protected natural areas do not become islands surrounded by development. In these situations, the site is effectively isolated, and its value for wildlife is greatly reduced. Careful planning can maintain natural environments along with the plants and animals associated with them. A balance between growth and the conservation of natural and scenic resources can be achieved by guiding development away from the most environmentally sensitive areas. The reclamation of previously disturbed areas for commercial and industrial projects, also known as brownfield development, presents one way to encourage economic growth while allowing ecologically sensitive areas to remain undisturbed. For example, reclaimed surface mines can be used for development (potentially even wind development) when feasible. Cluster development can be used to allow the same amount of development on much less land, and leave the remaining land intact for wildlife and native plants. By compressing development into already disturbed areas with existing infrastructure (villages, roads, existing rights-of-way), large pieces of the landscape can be maintained intact. If possible, networks or corridors of woodlands or greenspace should be preserved linking natural areas to each other. Preserving greenspace around development can provide ample recreation opportunities and potentially increase nearby property value. Manage for invasive species ­ Invasive species threaten native diversity by dominating habitat used by native species and by disrupting the integrity of the ecosystems they occupy. Small infestations may be easily controlled or eliminated, but larger, well established populations typically present difficult management challenges. The earlier exotic invasive species are identified and controlled, the greater the likelihood of eradication with the smallest expenditure of resources. Encourage conservation work by grassroots organizations ­ County and municipal governments can do much of the work necessary to plan for the protection and management of natural areas identified in this report; however, grassroots organizations are needed to assist with obtaining funding, identifying landowners who wish to protect their land, and providing information about easements, land acquisition, management, and stewardship of protected sites. Increasingly, local watershed organizations and land trusts are taking proactive steps to accomplish conservation at the local level. When activities threaten to impact ecological features, the responsible agency should be contacted. If no agency exists, private groups such as conservancies, land trusts, and watershed associations should be sought for ecological consultation and specific protection recommendations. Effects of Forest Roads on Macroinvertebrate Soil Fauna of the Southern Appalachian Mountains. Managing White-tailed Deer from an Ecosystem Perspective: Pennsylvania Case Study. Report by the Deer Management Forum for Pennsylvania Audubon and Pennsylvania Habitat Alliance, Harrisburg. Microenvironment Variation Within and Among Forest Edge Sites in the Eastern United States. Effects of Forest Fragmentation on Breeding Bird Populations in the Piedmont of the Mid-Atlantic Region. Pennsylvania Natural Heritage Program, Pennsylvania Department of Conservation and Natural Resources, Harrisburg, Pennsylvania. The reservoir was created in the early 1950s as a water source for several municipalities to the east. A small northeast-facing valley above the reservoir supports of population of the West Virginia white butterfly. This population may be unusual because it apparently depends on cut-leaved toothwort (Cardamine concatenata) as the host plant for its caterpillars. The most common host plant for this species in Pennsylvania is two-leaved toothwort (Cardamine diphylla), and there is evidence (Cappuchino and Kareiva, 1985) that cutleaved toothwort is not a reliable host plant because it dies back earlier than other toothworts, preventing caterpillars from reaching maturity. Seep scattered across the lower slopes of the site support a sensitive species of concern that is not named at request of the agency overseeing its protection. The sensitive species of concern depends on clean groundwater, and is vulnerable to changes in the quality or quantity of the groundwater. The result is that populations West Virginia white nectaring on yellow trout-lily (Erythronium americanum) are becoming genetically isolated, and if a population is extirpated the chances are low that remaining populations will be able to recolonize the habitat. An even greater threat to the West Virginia white is the spread of garlic mustard (Alliaria petiolata). This invasive plant is in the same family as toothworts (the mustard family), and the chemical signatures of the plants are similar enough that female butterflies will readily lay their eggs on garlic mustard. The caterpillars, however, cannot survive on garlic mustard, and these butterflies have disappeared from areas where garlic mustard is dominant. At this site garlic mustard is present at low densities, but widely distributed across the site. High densities of white-tailed deer pose another threat to West Virginia whites, because deer browsing greatly reduces the abundance of many of the wildflowers that are crucial nectar sources for the butterflies. The greatest threat to groundwater-dependent species like the sensitive species of concern is disruption to bedrock or any activity that alters groundwater flows or the quality of groundwater. High densities of water wells can depress water tables, causing springs and seeps to become dry or shrink in size. Groundwater pollution can occur from septic systems, improperly lined underground waste disposal, and, in agricultural areas, from infiltration of pesticides, fertilizers, or nutrients from animal wastes. A patch of lily-of-the-valley (Convallaria majalis) is near the remains of an old homestead.

Signature whistles of free-ranging bottlenose dolphins Tursiops truncatus: Stability and mother-offspring comparisons arthritis in dogs over the counter medication buy celebrex 200 mg online. Sex differences in signature whistle production in free-ranging bottlenose dolphins arthritis of neck exercises cheap celebrex 200 mg. Baseline arthritis in neck at 30 order celebrex 200 mg fast delivery, diurnal variations arthritis depression medication order celebrex 200 mg mastercard, and stress induced changes of stress hormones in three captive beluga whales arthritis in first joint of fingers buy celebrex 100mg otc, Delphinapterus leucas arthritis relief hands celebrex 200mg generic. A Discussion of Killer Whale Longevity (Orlando, Florida: SeaWorld Corporate Zoological Department). Psychological stress and the human immune system: A meta-analytic study of 30 years of inquiry. Age, growth, and maturity of bottlenosed dolphin (Tursiops truncatus) from Northeast Florida. The effects of human-dolphin interaction programmes on the behaviour of three captive Indo-Pacific humpback dolphins (Sousa chinensis). Review of the recent scientific data on the Okhotsk Sea white whale (Delphinapterus leucas) population structure and its application to management. Paper presented to the Scientific Committee at the 65th Meeting of the International Whaling Commission, 3­15 June 2013, Jeju Island, South Korea. Update report on the white whale (Delphinapterus leucas) live captures in the Okhotsk Sea, Russia. In Abstracts from the Ninth International Conference on Marine Mammals of the Holarctic (Astrakhan, Russia: Marine Mammal Council), p. Effects of pool size on free-choice selections by Atlantic bottlenose dolphins at one zoo facility. Thinking about Dolphins Thinking, Understanding the Impact of Social Narratives on Public Acceptance of Cognitive Science Research (New York. From captivity to the wild and back: An attempt to release Keiko the killer whale. Acclimation to captivity: A quantitative estimate based on survival of bottlenose dolphins and California sea lions. In vitro isolation and characterization of a calicivirus causing a vesicular disease of the hands and feet. Use of signature whistles during separations and reunions by wild bottlenose dolphin mothers and infants. Human neurobrucellosis with intracerebral granuloma caused by a marine mammal Brucella spp. Mission creep in the application of wildlife law: the progressive dilution of legal requirements regarding a wild-born orca kept for "research" purposes. Neuroimmunological response of beluga whales (Delphinapterus leucas) to translocation and a novel social environment. How do bottlenose dolphins, Tursiops truncatus, react to oil films under different light conditions? Dolphin thyroid and adrenal hormones: Circulating levels in wild and semi-domesticated Tursiops truncatus, and influence of sex, age, and season. Hematological, serum, and plasma chemical constituents in pantropical spotted dolphins (Stenella attenuata) following chase, encirclement, and tagging, Marine Mammal Science 29: 14­35. Polar Bears: the Natural History of a Threatened Species (Markham, Ontario: Fitzhenry & Whiteside). Immunologic evaluation of short-term captureassociated stress in free-ranging bottlenose dolphins (Tursiops truncatus) in Sarasota Bay. Paper presented at the 32nd Annual Conference of the International Association for Aquatic Medicine, Tampa, Florida, 28 April­2 May 2001. Paper presented at the 15th Biennial Conference on the Biology of Marine Mammals, Greensboro, North Carolina, 14­19 December 2003. New SeaWorld show just elephant drowning in large tank of water with no explanation. Free Willy: Phasing out captivity of killer whales with state level legislation and public support, Journal of Animal & Environmental Law 8: 22­23. Comparative analyses of longevity and senescence reveal variable survival benefits of living in zoos across mammals. Photo-identification catalogue and status of the northern resident killer whale population in 2014. Canadian Technical Report of Fisheries and Aquatic Sciences 3139 (Nanaimo, British Columbia: Department of Fisheries and Oceans), available at publications. The decline of Steller sea lions Eumetopias jubatus in Alaska: A review of the nutritional stress hypothesis. Does participation in dolphin-human interaction programs affect bottlenose dolphin behaviour? Baleen whale cortisol levels reveal a physiological response to 20th century whaling. Personality in captive killer whales (Orcinus orca): A rating approach based on the five-factor model. Journal of Comparative Psychology, advance online publication available at dx. Social and individual behavior of a group of bottlenose dolphins (Tursiops truncatus) in open and closed facilities. Behavior and salivary cortisol of captive dolphins (Tursiops truncatus) kept in open and closed facilities. An overview of increasing incidents of bottlenose dolphin harassment in the Gulf of Mexico and possible solutions. Driven by Demand: Dolphin Drive Hunts in Japan and the Involvement of the Aquarium Industry (Chippenham, United Kingdom: Whale and Dolphin Conservation Society). Emerging infectious diseases in cetaceans worldwide and the possible role of environmental stressors. Epidemiological pattern of tattoo skin disease: A potential general health indicator for cetaceans. Live-captures of common bottlenose dolphins Tursiops truncatus and unassessed bycatch in Cuban waters: Evidence of sustainability found wanting. Indeterminate status of West African populations of inshore common bottlenose dolphins Tursiops truncatus cautions against opportunistic live capture schemes. Primary bacterial pathogens in bottlenose dolphins Tursiops truncatus: Needles in haystacks of commensal and environmental microbes. Clinical relevance of urate nephrolithiasis in bottlenose dolphins, Tursiops truncatus. Evaluation of population health among bottlenose dolphins (Tursiops truncatus) at the United States Navy Marine Mammal Program. Hemochromatosis and fatty liver disease: Building evidence for insulin resistance in bottlenose dolphins (Tursiops truncatus). Blood-based indicators of insulin resistance and metabolic syndrome in bottlenose dolphins (Tursiops truncatus). Increased dietary intake of saturated fatty acid heptadecanoic acid (C17:0) associated with decreasing ferritin and alleviated metabolic syndrome in dolphins. Evaluation of annual survival and mortality rates and longevity of bottlenose dolphins (Tursiops truncatus) at the United States Navy Marine Mammal Program from 2004 through 2013. A Venezuelan court has ordered the start of trial against Waterland Mundo Marino Dolphinarium. Prolific body scars and collapsing dorsal fins on killer whales (Orcinus orca) in New Zealand waters. Ongoing Concerns with the SeaWorld Orca Held at Loro Parque, Tenerife, Spain (Unpublished report: Free Morgan Foundation), available at. Evaluating and minimising social stress in the care of captive bottlenose dolphins (Tursiops truncatus). Reassessing public opinion of captive cetacean attractions with a photo elicitation survey. SeaWorld emails show execs knew "Blackfish" hurt business long before they told investors. Personality and subjective well-being in orangutans (Pongo pygmaeus and Pongo abelii). The big none: No evidence for a general factor of personality in chimpanzees, orangutans, or rhesus macaques. Estimating bottlenose dolphin population parameters from individual identification and capture-release techniques. Evaluation of potential protective factors against metabolic syndrome in bottlenose dolphins: Feeding and activity patterns of dolphins in Sarasota Bay, Florida. Whale and Dolphin Conservation Society and the Humane Society of the United States (2003). Culture and conservation of non-humans with reference to whales and dolphins: Review and new directions. Beluga (Delphinapterus leucas) adult life expectancy: Wild populations vs the population in human care. In Georgia Aquarium (compiler), Application for a permit to import certain marine mammals for public display under the Marine Mammal Protection Act. A Comparison of Survival Rates for Free-Ranging Bottlenose Dolphins (Tursiops truncatus), Killer Whales (Orcinus orca), and Beluga Whales (Delphinapterus leucas). Urging cautious policy applications of captive research data is not the same as rejecting those data. Preliminary results about numerical discrimination in the bottlenose dolphin (Tursiops truncatus). Cultural transmission within maternal lineages: Vocal clans in resident killer whales in southern Alaska. Fatal necrotizing fasciitis and myositis in a captive common bottlenose dolphin (Tursiops truncatus) associated with Streptococcus agalactiae. Captive-born bottlenose dolphin x common dolphin (Tursiops truncatus x Delphinus capensis) intergeneric hybrids. Langseth in the Commonwealth of the Northern Mariana Islands, February­March 2012 Prepared for Lamont-Doherty Earth Observatory 61 Route 9W, P. The seismic study will use a towed array of 36 airguns with a total discharge volume of ~6600 in3. The proposed seismic survey is part of a research proposal recommended for funding by an expert review panel. It will provide data integral to advancing scientific understanding of the water cycle within subduction-zone systems, where the basic building blocks of continental crust are made and where Earth`s great earthquakes occur, which is thought to be the primary controlling factor in both arc-crust generation and megathrust seismicity. Potential impacts of the seismic survey on the environment would be primarily a result of the operation of the airgun array. Impacts would be associated with increased underwater noise, which may result in avoidance behavior by marine mammals, sea turtles, and fish, and other forms of disturbance. An integral part of the planned survey is a monitoring and mitigation program designed to minimize potential impacts of the proposed activities on marine animals present during the proposed research, and to document as much as possible the nature and extent of any effects. Injurious impacts to marine mammals, sea turtles, and seabirds have not been proven to occur near airgun arrays, and also are not likely to be caused by the other types of sound sources to be used. However, given the high levels of sound emitted by a large array of airguns, a precautionary approach is warranted. The planned monitoring and mitigation measures would reduce the possibility of injurious effects. The relatively wide shot spacing, in time and space, to be used during some of the survey, is an inherent mitigation measure relative to more typical seismic surveys with closer shotpoints. With the planned monitoring and mitigation measures, unavoidable impacts to each species of marine mammal and turtle that could be encountered are expected to be limited to short-term, localized changes in behavior and distribution near the seismic vessel. No long-term or significant effects are expected on individual marine mammals, sea turtles, seabirds, the populations to which they belong, or their habitats. Langseth meter Multibeam echosounder mile minute Protected Species Observer Protected Species Acoustic Observer (U. The proposed survey will provide data integral to advancing scientific understanding of the water cycle within subduction-zone systems, where the basic building blocks of continental crust are made and where Earth`s great earthquakes occur, which is thought to be the primary controlling factor in both arc-crust generation and megathrust seismicity. With these mitigation measures in place, any impacts on marine mammals and sea turtles are expected to be limited to short-term, localized changes in behavior of small numbers of animals. No long-term or significant effects are expected on individual mammals, turtles, or their populations. Impacts of seismic sounds on some pelagic seabirds are possible, although none are expected to be significant to individual birds or their populations. Subduction systems are where the basic building blocks of continental crust are made and where Earth`s great earthquakes occur. Little is known about either of these processes, but water cycling through the system is thought to be the primary controlling factor in both arc-crust generation and megathrust seismicity. An important new hypothesis has recently been suggested that, if correct, will transform our understanding of the water budget of subduction systems. This hypothesis holds that cracking attributable to bending of the subducting plate enables water to penetrate through the subducting crust into the mantle, where it hydrates the mantle by forming the hydrous mineral phase serpentine. This phase is stable to greater depths than the hydrous clay minerals of the crust, where most of the subducting water was previously believed to be held. Thus, if this hypothesis is correct, it provides a mechanism for transporting water far beneath the mantle wedge, where it promotes melting and crust formation, and possibly even deeper into the mantle, providing a whole-earth hydration mechanism that promotes the continued operation of plate tectonics, without which our planet would likely be unable to support life. The scientists involved in this program will test this hypothesis by measuring mantle seismic sounds speeds, which vary with degree of serpentinization. By comparing these measurements from the Mariana system, which is old and cold, with the Costa Rica system, which is young and warm and where similar measurements have recently been made, we should be able to definitively determine whether or not substantial water is taken up by the mantle of subducting plates near the outer rise of seafloor trenches. The closest that the vessel will approach to any island is ~50 km from Alamagan. As the airgun array is towed along the survey lines, the hydrophone streamer will receive the returning acoustic signals and transfer the data to the on-board processing system. This includes one line and parts of three lines shown in Figure 1 that are shot twice at different shot intervals: the westernmost north-south line and the western portions of the three east-west lines.

In the Southern Hemisphere arthritis relief using gelatin cheap celebrex 200mg fast delivery, mating occurs from July to March arthritis lower back causing leg pain cheap celebrex 100 mg otc, with a peak from September to December rheumatoid arthritis remission purchase 100mg celebrex with mastercard, and most calves are born between November and March (Rice 1989) arthritis diet wine generic 100mg celebrex. Sperm whales generally are distributed over large areas that have high secondary productivity and steep underwater topography rheumatoid arthritis nails purchase celebrex 200 mg free shipping, in waters at least 1000 m deep (Jaquet and Whitehead 1996; Whitehead 2002a) arthritis joint protection handout celebrex 200 mg mastercard. They are often found far from shore, but can be found closer to oceanic islands that rise steeply from deep ocean waters (Whitehead 2002a). They can dive as deep as ~2 km and possibly deeper on rare occasions for periods of over 1 h; however, most of their foraging occurs at depths of ~300­800 m for 30­ 45 min (Whitehead 2003). Whales in the Galбpagos Islands typically dove for ~40 min and then spent 10 min at the surface (Papastavrou et al. Whaling records are consistent with sperm whales occurring year-round around the Marianas (Townsend 1935), and there are two stranding records for Guam (Kami and Lujan 1976; Eldredge 1991, 2003). Sperm whales were observed in waters ~800 to 10,000 m deep throughout most of the survey area. Observations were made during the January­April 2007 survey of several large bulls with fresh tooth marks (one male rammed the survey ship), which suggests that these males were engaged in competition for mates (Fulling and Salinas Vega 2009). Hence, there is evidence that this area is used for breeding and calving by sperm whales. They are difficult to sight at sea, perhaps because of their avoidance reactions to ships and behavior changes in relation to survey aircraft (Wьrsig et al. The two species are difficult to distinguish from one another when sighted (McAlpine 2002). During sighting surveys and, hence, in population and density estimates, the two species are most often categorized together as Kogia spp. Affected Environment Pygmy sperm whales may inhabit waters beyond the continental shelf edge, whereas dwarf sperm whales are thought to inhabit the shelf-edge and slope waters (Rice 1998). Also, the dwarf sperm whale could prefer warmer waters than the pygmy sperm whale (McAlpine 2002). Pygmy sperm whales feed mainly on various species of squid in the deep zones of the continental shelf and slope (McAlpine et al. Pygmy sperm whales occur in small groups of up to six, and dwarf sperm whales can form groups of up to 10 (Caldwell and Caldwell 1989). Although there are few useful estimates of abundance for pygmy or dwarf sperm whales anywhere in their range, they are thought to be fairly common in some areas. Its inconspicuous blows, deepdiving behavior, and tendency to avoid vessels all help to explain the infrequent sightings (Barlow and Gisner 2006). Adult males of this species usually travel alone, but these whales can be seen in groups of up to 15 individuals, with a mean group size of 2. Cuvier`s beaked whale is an offshore, deep-diving species that feeds on fish and squid (Heyning 2002). Its dives generally last 30­60 min, but dives of 85 min have been recorded (Tyack et al. In the western Pacific, Cuvier`s beaked whales are known to occur in the waters of Japan (Nishiwaki and Oguro 1972 in Wang et al. Recent morphometric and genetic analyses of those two original specimens and an additional four specimens have allowed a more detailed characterization of the species (Dalebout et al. It seems likely that it is, in fact, the cetacean that has been seen in IndoPacific waters and called the tropical bottlenose whale. Some authorities place the species in the genus Mesoplodon, but there now seems to be sufficient information to afford it status as a separate genus (Dalebout et al. Longman`s beaked whales have been sighted in waters with temperatures 21­31єC and have been seen in the tropics every month of the year except June, indicating year-round residency (Pitman et al. Although widespread throughout the tropical Pacific, the species must still be considered rare because of a scarcity of sightings despite a great deal of survey effort (Pitman et al. Longman`s beaked whales have been seen alone, but more commonly in groups of at least ten and up to 100, with an average group size of 15­20 (Jefferson et al. Sightings of Longman`s beaked whale have occurred at many locations in tropical waters of the Indo-Pacific region (Rudolph and Smeenk 2002; Jefferson et al. Mesoplodont Beaked Whales Two species of mesoplodont whales likely occur in deep waters in the study area: Blainville`s beaked whale (Mesoplodon densirostris) and the gingko-toothed beaked whale (Mesoplodon ginkgodens). Almost everything that is known regarding most mesoplodont species has come from stranded animals (Pitman 2002). The different mesoplodont species are difficult to distinguish in the field, and are most often categorized during sighting surveys, and therefore in density and population estimates, as Mesoplodon spp. They are all thought to be deep-water animals, only rarely seen over the continental shelf. Blainville`s beaked whale has the widest distribution throughout the world of all Mesoplodon species (Mead 1989). Blainville`s beaked whales are most often found in singles or pairs, but also in groups of 3­7 (Jefferson et al. Like other beaked whales, Blainville`s beaked whales are generally found in deep waters 200 m to 1400 m deep (Gannier 2000; Jefferson et al. However, they also can occur in coastal areas and have been known to spend long periods of time at depths <50 m (Jefferson et al. Sighting records exist for Blainville`s beaked whale for the East China Sea off mainland China and for the Philippines (Perrin et al. In the South Pacific Ocean, it has stranded in New South Wales, Australia, and the North Island and Chatham Islands, New Zealand (Mead 1989; Baker and van Helden 1999). Rough-toothed Dolphin (Steno bredanensis) the rough-toothed dolphin is widely distributed around the world, but mainly occurs in tropical and warm temperate waters (Miyazaki and Perrin 1994). Rough-toothed dolphins generally occur in deep, oceanic waters, but can be found in shallower coastal waters in some regions (Jefferson et al. Rough-toothed dolphins are deep divers and can dive for up to 15 min (Jefferson et al. They usually form groups of 10­20, but aggregations of hundreds have been seen (Jefferson et al. Rough-toothed dolphins are known to occur in the Philippines and the East and South China seas off China and Taiwan (Perrin et al. The sightings were in deep (1000­4500 m) water, one off Guguan and the other at the southern edge of the survey area at ~10єN. One sighting involved a mixed-species group with common bottlenose dolphins and short finned pilot whales. In July 2004, there was a sighting of an undetermined smaller number of rough-toothed dolphins mixed in with a school of an estimated 500­700 melon-headed whales at Sasanhayan Bay, Rota, in a water depth of ~75 m (Jefferson et al. Common Bottlenose Dolphin (Tursiops truncatus) the bottlenose dolphin is distributed worldwide. Generally, there are two distinct bottlenose dolphin types: a shallow water type, mainly found in coastal waters, and a deep water type, mainly found in oceanic waters (Duffield et al. As well as inhabiting different areas, these ecotypes differ in their diving abilities (Klatsky 2004) and prey types (Mead and Potter 1995). Bottlenose dolphins have been reported to regularly dive to depths >450 m for periods of >5 min (Klatsky 2004), and even down to depths of 600­700 m for up to 12 min (Klatsky et al. In the western Pacific, the bottlenose dolphin is distributed from Japan to Australia and New Zealand. Bottlenose dolphins are known to occur in the Philippines and the Yellow, East and South China seas off China and Taiwan (Perrin et al. Two of the sightings were in the vicinity of Challenger Deep at the southwest corner of the 2007 survey area, and the other was east of Saipan near the Mariana Trench in water depths 4200­5000 m. One of the sightings near the Challenger Deep was a mixed-species aggregation that included sperm whales (with calves) logging at the surface. Another mixed-species aggregation involved bottlenose dolphins with short-finned pilot whales and rough-toothed dolphins. There are no stranding records available for this species in the Marianas, and only a mention by Trianni and Kessler (2002) that bottlenose dolphins are seen in coastal waters of Guam. It is possible that bottlenose dolphins do not occur in great numbers in this island chain. Affected Environment that large densities of bottlenose dolphins do not occur at the Marquesas Islands to the fact that the area does not have a significant shelf component. Pantropical Spotted Dolphin (Stenella attenuata) the pantropical spotted dolphin can be found throughout tropical and some subtropical oceans of the world (Perrin and Hohn 1994). The offshore form inhabits tropical, equatorial, and southern subtropical water masses (Perrin 2002a). This species is found primarily in deeper waters, and rarely over the continental shelf or continental shelf edge (Davis et al. Pantropical spotted dolphins are extremely gregarious, forming groups of hundreds or even thousands. Pantropical spotted and spinner dolphins are commonly seen together in mixed-species groups. In the western Pacific, pantropical spotted dolphins occur from Japan south to Australia. They are known to occur in the Philippines and in the East and South China seas off China (Perrin et al. Most (11 of 17) sightings were in deep (>3000 m) water, which is consistent with the known preference of this species for oceanic habitat. Group size was 1­115, and there were multiple sightings that included calves, one mixed-species aggregation with melon-headed whales, and another with an unidentified rorqual. Spinner Dolphin (Stenella longirostris) the spinner dolphin is distributed in oceanic and coastal tropical waters between 40єN and 40єS (Jefferson et al. Two subspecies of spinner dolphin occur in the western Pacific: the widespread, offshore spinner dolphin (Stenella longirostris longirostris) and the dwarf spinner dolphin (S. There is little or no genetic interchange between the two subspecies (Dizon et al. The spinner dolphin is expected to occur throughout the Mariana Islands, except in Apra Harbor because of likely avoidance of anthropogenic disturbances there (DoN 2005). Spinner dolphins are reported in Saipan Lagoon nearly every year (Trianni and Kessler 2002). Affected Environment will congregate in lagoons during the day to rest and avoid predation. Spinner dolphins travel among the Mariana island chain (Trianni and Kessler 2002) and have been seen at Farallon de Medinilla (DoN 2001; Trianni and Kessler 2002), Guam (Trianni and Kessler 2002), and Rota (Jefferson et al. The school was encountered northeast of Saipan in 425 m of water, and was estimated to contain 98 individuals. Striped Dolphin (Stenella coeruleoalba) the striped dolphin has a cosmopolitan distribution in tropical to warm temperate waters (Perrin et al. It is typically found in waters outside the continental shelf and is often associated with convergence zones and areas of upwelling (Archer 2002). Striped dolphins are fairly gregarious (groups of 20 or more are common) and active at the surface (Whitehead et al. A putative population south of 30°N in the western Pacific has been estimated to number about 52,600 (Miyashita 1993), and any animals around the Mariana Islands are probably part of this greater population. Prior to 2007, striped dolphins were only known from one stranding that occurred in July 1985 (Wilson et al. It only occurs rarely in temperate regions, and then only in relation to temporary oceanographic anomalies such as El Niсo events (Perrin et al. Off Huahine and Tahiti (Society Islands), it was observed in waters 500­1500 m deep (Gannier 2000). Fraser`s dolphins travel in groups ranging from just a few animals to 100 or even 1000 (Perrin et al. In the eastern Sulu Sea adjacent to the Philippines, the abundance estimate is 8700 (Dolar 1999 in Perrin et al. However, the Mariana Islands are within the distributional range of this species (DoN 2005). Short-beaked Common Dolphin (Delphinus delphis) the common dolphin is found in tropical and warm temperate oceans around the world (Perrin 2002b). It ranges as far south as 40°S in the Pacific Ocean, is common in coastal waters 200­300 m deep, and is also associated with prominent underwater topography, such as seamounts (Evans 1994). Common dolphins often travel in fairly large groups; schools of hundreds or even thousands are common. Smith and Whitehead (1999) noted that common dolphins were frequently seen in waters near the Galбpagos Islands, with a mean group size of 125. The short-beaked common dolphin is the species most likely to occur in the Mariana Islands (Jefferson pers. It occurs between 60єN and 60єS, where surface water temperatures are at least 10єC (Kruse et al. In the northern Gulf of Mexico, Risso`s dolphin usually occurs over steeper sections of the upper continental slope (Baumgartner 1997) in waters 150­2000 m deep (Davis et al. In Monterey Bay, California, it is most numerous where there is steep bottom topography (Kruse et al. Risso`s dolphins occur individually or in small to moderate-sized groups, normally ranging from 2 to <250. In the western Pacific, Risso`s dolphins range from the Kuril Islands to New Zealand and Australia. They are known to occur in the Philippines, off mainland China in the Yellow, East, and South China seas (Perrin et al. Under the assumption that several stocks exist in the western Pacific, Miyashita (1993) analyzed Japanese survey data to estimate that about 7000 Risso`s dolphins occur in the area to the north of the Mariana Islands. Melon-headed Whale (Peponocephala electra) the melon-headed whale is a pantropical and pelagic species that occurs mainly between 20єN and 20єS in offshore waters (Perryman et al. Melon-headed whales tend to occur in groups of 100­ 500, but have also been seen in groups of up to 2000 (Jefferson et al. Melon-head whales are commonly seen in mixed groups with other cetaceans (Jefferson and Barros 1997).

O ­ Proline Biosynthesis of proline from glutamate employs reactions similar to those of proline catabolism arthritis pain relief jewelry cheap 100mg celebrex otc, but in which glutamate -phosphate is an intermediate (Figure 27­8) arthritis pain apply heat or cold celebrex 100 mg amex. Following conversion of methionine to homocysteine (see Figure 29­19) arthritis flare up in neck buy celebrex 100mg free shipping, homocysteine and serine form cystathionine arthritis pain doterra buy cheap celebrex 100 mg, whose hydrolysis forms cysteine and homoserine (Figure 27­9) arthritis knee grade 3 generic 200 mg celebrex with visa. Note similarities to and differences from the glutamine synthetase reaction (Figure 27­2) arthritis toes celebrex 100mg with mastercard. The enzymes that catalyze the reactions shown are choline dehydrogenase, betaine dehydrogenase, betaine-homocysteine N-methyltransferase, sarcosine demethylase, and sarcosine oxidase, respectively. The catalysts for these reactions are glutamate 5-kinase, glutamate semialdehyde dehydrogenase, noncatalyzed ring closure, and pyrolline 5-carboxylate reductase. If the diet contains adequate quantities of the nutritionally essential amino acid phenylalanine, tyrosine is nutritionally nonessential. But since the phenylalanine hydroxylase reaction is irreversible, dietary tyrosine cannot replace phenylalanine. Catalysis by this mixed-function oxygenase incorporates one atom of O2 into the para position of phenylalanine and reduces the other atom to water. The sulfur of cysteine derives from methionine and the carbon skeleton from serine. This reaction is associated with several defects of phenylalanine metabolism discussed in Chapter 29. During the course of the reaction, molecular oxygen is incorporated into both succinate and proline. Unlike hydroxyproline or hydroxylysine, selenocysteine arises co-translationally during its incorporation into peptides. Subsequent replacement of the serine oxygen by selenium involves selenophosphate formed by selenophosphate synthase (Figure 27­12). Selenocysteine (top) and the reaction catalyzed by selenophosphate synthetase (bottom). Hydroxylation of peptidyl prolyl and lysyl residues, catalyzed by prolyl hydroxylase and lysyl hydroxylase of skin, skeletal muscle, and granulating wounds requires, in addition to the substrate, molecular O2, ascorbate, Fe2+, and -ketoglutarate (Figure 27­11). For every mole of proline or lysine hydroxylated, one mole of -ketoglutarate is decarboxylated to succinate. One atom of O2 is incorporated into proline or lysine, the other into succinate (Figure 27­11). The intermediates and the amino acids to which they give rise are -ketoglutarate (Glu, Gln, Pro, Hyp), oxaloacetate (Asp, Asn), and 3-phosphoglycerate (Ser, Gly). Cysteine, tyrosine, and hydroxylysine are formed from nutritionally essential amino acids. Serine provides the carbon skeleton and homocysteine the sulfur for cysteine biosynthesis. Phenylalanine hydroxylase converts phenylalanine to tyrosine in an irreversible reaction. Peptidyl hydroxyproline and hydroxylysine are formed by hydroxylation of peptidyl proline or lysine in reactions catalyzed by mixed-function oxidases that require vitamin C as cofactor. The nutritional disease scurvy reflects impaired hydroxylation due to a deficiency of vitamin C. Selenocysteine, the 21st Amino Acid While its occurrence in proteins is uncommon, selenocysteine(Figure 27­12) is present at the active site of several human enzymes that catalyze redox reactions. Examples include thioredoxin reductase, glutathione peroxidase, and the deiodinase that converts thyroxine to triiodothyronine. Where present, selenocysteine participates in the catalytic mechanism of these enzymes. Significantly, replacement of selenocysteine by cysteine can significantly decrease catalytic activity. Impairments in human selenoproteins have been implicated in tumorigenesis and atherosclerosis, and are associated with selenium deficiency cardiomyopathy (Keshan disease). Stickel F et al: Role of nutrition in liver transplantation for end-stage chronic liver disease. Positive nitrogen balance, an excess of ingested over excreted nitrogen, accompanies growth and pregnancy. Negative nitrogen balance, where output exceeds intake, may follow surgery, advanced cancer, kwashiorkor, and marasmus. Tissues therefore convert ammonia to the amide nitrogen of the nontoxic amino acid glutamine. Subsequent deamination of glutamine in the liver releases ammonia, which is then converted to urea, which is not toxic. If liver function is compromised, as in cirrhosis or hepatitis, elevated blood ammonia levels generate clinical signs and symptoms. Each enzyme of the urea cycle provides examples of metabolic defects and their physiologic consequences, and the cycle as a whole serves as a molecular model for the study of human metabolic defects. Each day, humans turn over 1­2% of their total body protein, principally muscle protein. High rates of protein degradation occur in tissues that are undergoing structural rearrangement, for example, uterine tissue during pregnancy, skeletal muscle in starvation, and tadpole tail tissue during metamorphosis. Since excess free amino acids are not stored, those not immediately incorporated into new protein are rapidly degraded. The major portion of the carbon skeletons of the amino acids is converted to amphibolic intermediates, while the amino nitrogen is converted to urea and excreted in the urine. The resulting peptides are then degraded to amino acids by endopeptidases that cleave internal peptide bonds, and by aminopeptidases and carboxypeptidases that remove amino acids sequentially from the amino and carboxyl terminals, respectively. Degradation of circulating peptides such as hormones follows loss of a sialic acid moiety from the nonreducing ends of their oligosaccharide chains. Asialoglycoproteins are internalized by liver cell asialoglycoprotein receptors and degraded by lysosomal proteases termed cathepsins. Ubiquitin molecules are attached by non-peptide bonds formed between the carboxyl terminal of ubiquitin and the -amino groups of lysyl residues in the target protein (Figure 28­1). The residue present at its amino terminal affects whether a protein is ubiquitinated. Attachment of a single ubiquitin molecule to transmembrane proteins alters their subcellular localization and targets them for degradation. Soluble proteins undergo polyubiquitination, further ligase-catalyzed attachment of four or more additional ubiquitin molecules. Subsequent degradation of ubiquitin-tagged proteins takes place in the proteasome, a macromolecule with multiple different subunits that also is ubiquitous in eukaryotic cells. The key role of alanine in amino acid output from muscle and gut and uptake by the liver is shown. Alanine, which appears to be the vehicle of nitrogen transport in the plasma, is extracted primarily by the liver. Glutamine is extracted by the gut and the kidney, both of which convert a significant portion to alanine. The kidney provides a major source of serine for uptake by peripheral tissues, including liver and muscle. Branched-chain amino acids, particularly valine, are released by muscle and taken up predominantly by the brain. The rate of hepatic gluconeogenesis from alanine is far higher than from all other amino acids. The capacity of the liver for gluconeogenesis from alanine does not reach saturation until the alanine concentration reaches 20­30 times its physiologic level. Following a protein-rich meal, the splanchnic tissues release amino acids (Figure 28­4) while the peripheral muscles extract amino acids, in both instances predominantly branched-chain amino acids. Branched-chain amino acids thus serve a special role in nitrogen metabolism, both in the fasting state, when they provide the brain with an energy source, and after feeding, when they are extracted predominantly by muscles, having been spared by the liver. While depicted as single entities, there are several types of E1, and over 500 types of E2. E3 then catalyzes the transfer of ubiquitin to the -amino group of a lysyl residue of the target protein. Metabolic diseases associated with defects of ubiquitination include the Angelman syndrome and the von Hippel-Lindau syndrome in which there is a defect in the ubiquitin E3 ligase. For additional aspects of protein degradation and of ubiquitination, including its role in the cell cycle, see Chapter 4. Muscle generates over half of the total body pool of free amino acids, and liver is the site of the urea cycle enzymes necessary for disposal of excess nitrogen. Muscle and liver thus play major roles in maintaining circulating amino acid levels. The aqueous environment of teleostean fish, which are ammonotelic (excrete ammonia), compels them to excrete water continuously to facilitate excretion of the highly toxic molecule ammonia. Birds, which must conserve water and maintain low weight, are uricotelic and excrete uric acid as semisolid guano. Many land animals, including humans, are ureotelic and excrete nontoxic, water-soluble urea. Of the protein amino acids, all except lysine, threonine, proline, and hydroxyproline participate in transamination. Rearrangement forms an -keto acid and enzyme-bound pyridoxamine phosphate, which then forms a Schiff base with a second keto acid. Following removal of its -amino nitrogen by transamination, the remaining carbon "skeleton" of an amino acid is degraded by pathways discussed in Chapter 29. Alanine-pyruvate aminotransferase (alanine aminotransferase) and glutamate-ketoglutarate aminotransferase (glutamate aminotransferase) catalyze the transfer of amino groups to pyruvate (forming alanine) or to -ketoglutarate (forming glutamate) (Figure 28­7). Alanine is synthesized in muscle by transamination of glucose-derived pyruvate, released into the bloodstream, and taken up by the liver. In the liver, the carbon skeleton of alanine is reconverted to glucose and released into the bloodstream, where it is available for uptake by muscle and resynthesis of alanine. H2O2 O2 O R C C O -Keto acid O­ is specific for one pair of substrates, but nonspecific for the other pair. Since alanine is also a substrate for glutamate aminotransferase, all the amino nitrogen from amino acids that undergo transamination can be concentrated in glutamate. This is important because l-glutamate is the only amino acid that undergoes oxidative deamination at an appreciable rate in mammalian tissues. The formation of ammonia from -amino groups thus occurs mainly via the -amino nitrogen of l-glutamate. The -amino group of ornithine (but not the -amino group of lysine) readily undergoes transamination. Elevated serum levels of aminotransferases characterize certain diseases (see Table 7­2). The reduced flavin is reoxidized by molecular oxygen, forming hydrogen peroxide (H2O2), which then is split to O2 and H2O by catalase. Ammonia Intoxication Is Life-Threatening the ammonia produced by enteric bacteria and absorbed into portal venous blood and the ammonia produced by tissues are rapidly removed from circulation by the liver and converted to urea. Should portal blood bypass the liver, systemic blood ammonia levels may attain toxic levels. This occurs in severely impaired hepatic function or the development of collateral links between the portal and systemic veins in cirrhosis. Symptoms of ammonia intoxication include tremor, slurred speech, blurred vision, coma, and ultimately death. Ammonia may be toxic to the brain in part because it reacts with -ketoglutarate to form glutamate. Amino Acid Oxidases Also Remove Nitrogen as Ammonia While their physiologic importance is uncertain, l-amino acid oxidases of liver and kidney convert an amino acid to an -imino acid that decomposes to an -keto acid with re- Glutamine Synthetase Fixes Ammonia as Glutamine Formation of glutamine is catalyzed by mitochondrial glutamine synthetase (Figure 28­10). Glutaminase & Asparaginase Deamidate Glutamine & Asparagine Glutamine synthetase plays a major role in ammonia detoxification, interorgan nitrogen flux, and acid-base homeostasis. Of the six participating amino acids, N-acetylglutamate functions solely as an enzyme activator. The major metabolic role of ornithine, citrulline, and argininosuccinate in mammals is urea synthesis. The ornithine consumed in reaction 2 is regenerated in reaction 5, and so there is no net loss or gain of ornithine, citrulline, argininosuccinate, or arginine. Some reactions of urea synthesis occur in the matrix of the mitochondrion, and other reactions in the cytosol (Figure 28­12). The concerted action of glutamine synthetase and glutaminase thus catalyzes the interconversion of free ammonium ion and glutamine. A rare deficiency in neonate glutamine synthetase results in severe brain damage, multi-organ failure, and death. Carbamoyl Phosphate Plus Ornithine Forms Citrulline l-Ornithine transcarbamoylase catalyzes transfer of the carbamoyl group of carbamoyl phosphate to ornithine, forming citrulline and orthophosphate (reaction 2, Figure 28­12). While the reaction occurs in the mitochondrial matrix, both the formation of ornithine and the subsequent metabolism of citrulline take place in the cytosol. Entry of ornithine into mitochondria and exodus of citrulline from mitochondria therefore involve mitochondrial inner membrane transport systems (Figure 28­12). Formation & Secretion of Ammonia Maintains Acid-Base Balance Excretion into urine of ammonia produced by renal tubular cells facilitates cation conservation and regulation of acid-base balance. Ammonia production from intracellular renal amino acids, especially glutamine, increases in metabolic acidosis and decreases in metabolic alkalosis. The nitrogen-containing groups that contribute to the formation of urea are shaded. Reactions and occur in the matrix of liver mitochondria and reactions, and in liver cytosol. Citrulline Plus Aspartate Forms Argininosuccinate Argininosuccinate synthetase links aspartate and citrulline via the amino group of aspartate (reaction 3, Figure 28­12) and provides the second nitrogen of urea.

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