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Sensory ability in the narwhal tooth organ system. Correspondence to: Martin T. Nweeia, Harvard School of essays charles Dental Medicine, Department of Restorative Dentistry and essays Biomaterial Sciences, 188 Longwood Ave., Boston, MA. E-mail: Martin_nweeia@hsdm.harvard.edu. Delta Dental of Wisconsin, P.O. Box 828, Stevens Point, WI Search for more papers by charles this author. Orthopaedic Research Center, Boston Children's Hospital, 300 Longwood Ave., Boston, MA Department of Orthopaedic Surgery, Harvard Medical School, 25 Shattuck Street, Boston, MA Department of why i response Developmental Biology, Harvard School of Dental Medicine, 188 Longwood Ave., Boston, MA Search for more papers by this author. Carlson School of Management, University of essays lamb Minnesota, 321 19th Ave. S., Minneapolis, MN Search for more papers by this author. Arctic Research Division, Fisheries and water Oceans Canada, 501 University Crescent, Winnipeg, MB, Canada Search for more papers by english charles this author. Freshwater Institute, Fisheries and Oceans Canada, 501 University Crescent, Winnipeg, Canada Search for answers more papers by this author. Freshwater Institute, Fisheries and Oceans Canada, 501 University Crescent, Winnipeg, Canada Search for more papers by this author.
Department of Vertebrate Zoology, Smithsonian Institution, 1000 Jefferson Drive SW, Washington, DC Search for more papers by this author. Department of english charles Vertebrate Zoology, Smithsonian Institution, 1000 Jefferson Drive SW, Washington, DC Search for more papers by narrative essay this author. Arctic Research Center, Institute of Bioscience, Aarhus University, Frederiksborgvej, Roskilde, Denmark Search for more papers by english essays lamb this author. National Institute of Standards and Technology, ADAF Paffenbarger Research Center, 100 Bureau Drive, Gaithersburg, MD Search for more papers by this author. Veterinary Services, Calgary Zoo, 1300 Zoo Rd. NE, Calgary, AB, Canada Search for more papers by this author. Catalyst Laboratory for Innovative Translational Technologies, Harvard School of Dental Medicine, 188 Longwood Ave., Boston, MA Search for more papers by this author. Department of Developmental Biology, Harvard School of Dental Medicine, 188 Longwood Ave., Boston, MA Catalyst Laboratory for Innovative Translational Technologies, Harvard School of essay Dental Medicine, 188 Longwood Ave., Boston, MA Search for more papers by this author. First published: 18 March 2014 Full publication history DOI: 10.1002/ar.22886 View/save citation Cited by (CrossRef): 6 articles Check for english essays charles updates.
Animal Care : Animal Protocol, 04384. Short Format Answers? Office of Research Subject Protection, Harvard Medical School. The erupted tusk of the narwhal exhibits sensory ability. The hypothesized sensory pathway begins with ocean water entering through cementum channels to a network of patent dentinal tubules extending from the dentinocementum junction to the inner pulpal wall. English Lamb? Circumpulpal sensory structures then signal pulpal nerves terminating near the friendships, base of the tusk. English Essays Charles Lamb? The maxillary division of the fifth cranial nerve then transmits this sensory information to kids, the brain. This sensory pathway was first described in published results of patent dentinal tubules, and evidence from english lamb, dissection of tusk nerve connection via the water kids, maxillary division of the fifth cranial nerve to the brain. New evidence presented here indicates that the patent dentinal tubules communicate with open channels through a porous cementum from the ocean environment. The ability of pulpal tissue to english essays charles lamb, react to external stimuli is supported by immunohistochemical detection of neuronal markers in the pulp and gene expression of pulpal sensory nerve tissue. Final confirmation of sensory ability is water, demonstrated by essays charles significant changes in heart rate when alternating solutions of high-salt and essay in life fresh water are exposed to the external tusk surface.
Additional supporting information for function includes new observations of dentinal tubule networks evident in unerupted tusks, female erupted tusks, and essays vestigial teeth. New findings of sexual foraging divergence documented by stable isotope and fatty acid results add to the discussion of the narrative essay, functional significance of the narwhal tusk. The combined evidence suggests multiple tusk functions may have driven the tooth organ system's evolutionary development and persistence. Anat Rec, 297:599617, 2014. English Charles? 2014 Wiley Periodicals, Inc.
The functional significance of the erupted narwhal tusk has been the subject of word essay on accountability and responsibility conjecture and charles theory since the christian, writings of Albertus Magnus in english charles lamb, 1495 (Magnus, 1495). It has been thought to serve as an acoustic probe (Best, 1981; Reeves and Mitchell, 1981), possibly associated with sound transmission (Ford and Fisher, 1978; Best, unpublished B.Sc. Thesis, University of British Columbia, 1972); a thermal regulator (Dow and Hollenberg, 1977); a swimming rudder (Kingsley and Ramsay, 1988); a breathing organ; a spear for essays kids hunting or finding food (Vibe, 1950; Harrison and King, 1965; Ellis, 1980; Bruemmer, 1993); an essays charles, aggressive weapon in interspecific fighting (Brown, 1868; Beddard, 1900; Lowe, 1906; Geist et al., 1960) or self-defense against predators (Buckland, 1882; Gray, 1889: Freuchen, 1935); and a tool used for breaking the ice (Scoresby, 1820; Tomlin, 1967), digging (Freuchen, 1935; Pederson, 1960; Newman, 1971), or resting on narrative, ice (Porsild, 1918). Many studies describe the tusk as a secondary sexual characteristic used in aggressive encounters or intraspecific display (Lowe, 1906; Norman and english essays Fraser, 1949; Miller, 1955; Silverman, 1979) associated with tusk fracture and head scarring (Porsild, 1922; Silverman and Dunbar, 1980), and to establish social hierarchy amongst males (Scoresby, 1820; Hartwig, 1874; Mansfield et al., 1975; Silverman and Dunbar, 1980; Gerson and Hickie, 1985). Examining the essay answers, narwhal tooth organ system through a multidisciplinary approach that combines studies of anatomy, morphology, histology, neurophysiology, genetics, and diet gives a more comprehensive view of its functional significance and highlights sensory ability as an added functional attribute. Narwhal teeth have unusual anatomical features. Among them are (1) a sinistral spiral morphology (Worm, 1655; Scoresby, 1823); (2) an extreme degree of tooth asymmetry in males, with a single left tusk expression and embedded right tusk (Sonnini and english lamb Buffon, 1804; Home, 1813); (3) an extreme expression of sexual dimorphism, with the 1000 word essay, male having an erupted left canine tusk reaching 2.6 m and the female commonly with right and left embedded tusks, often less than 33 cm (Sonnini and Buffon, 1804; Home, 1813); (4) a unique form of tooth asymmetry in lamb, a double-tusked expression, since the morphology of the narrative essay, spiral remains sinistral for essays charles lamb both left and essay right antemeres, and the left tusk is often slightly longer than the right (Worm, 1655; Sonnini and Buffon, 1804; Home, 1813); (5) a horizontal direction of eruption in both erupted and english unerupted tusk forms (Linné et al., 1792: Egede and Wood, 1818); and (6) perforation of the tooth through the on regrets in life, upper lip (Hampe, 1737; Brisson, 1756; Crantz, 1767; Donndorff, 1792).
Though narwhal teeth share many anatomical characteristics with other tusked animals including a lack of enamel (Ishiyama, 1987), the presence of essays charles lamb cementum, dentin, pulpal tissue and their associated structures (Seltzer and Bender, 2002; Berkovitz et al., 2002), and the presence of the want a wife response, maxillary division of the fifth cranial nerve associated with tooth innervation (Nweeia et al., 2009), they have many distinguishing features. Among them are a cementum layer overlying over the erupted canine (Nweeia et al., 2012) tusk, a patent network of dentinal tubules through the full thickness of dentin (Nweeia et al., 2009; Boyde, 1980; Locke, 2008), and pulpal soft tissues, extending the full length of the tooth, diminishing only in diameter with age (Pederson, 1931; Dow and english essays Hollenberg, 1977; Nweeia et al., 2009). Initial scanning electron micrographs of the erupted male narwhal tusk reveal patent dentinal tubules that extend the full thickness of the essay, dentin and correspond to lumina on the tusk surface. The tubules radiating outward from the english charles, dentin-pulpal wall are similar to those of humans and other mammals in diameter, though the spacing is three to five times wider. Limited scanning electron microscopy (SEM) has been completed for other odontocetes, but suggests dentinal tubules are well occluded within the erupted portion of the dentinal layer for most odontocetes (Boyde, 1980). In cases where the dentinal tubules are patent through the full dentinal layer, such as the sperm whale ( Physeter macrocephalus ) (Boyde, 1980; Locke, 2008), they are covered by an enamel layer (Loch et al., 2012). Dentinal tubules have been described for other marine mammals, including the essay, rough-toothed dolphin ( Steno bredanensis ) (Miyazaki, 1977), pantropical spotted dolphin ( Stenella attenuata ) (Myrick, 1980), grey seal ( Halichoerus grypus ) (Hewer, 1964), hooded seal ( Cystophora cristata ) (Mohr, 1966) harbor porpoise ( Phoecoena phoecoena ) (Perrin and Myrick, 1980), short-beaked dolphin ( Delphinus delphis ) (Gurevich et al., 1980), bottlenose dolphin ( Tursiops truncatus ), and pilot whale ( Globicephala melaena ) (Boyde, 1980; Locke, 2008), and essays lamb the walrus ( Odobenus rosmarus ), hippopotamus ( Hippopotamus amphibious ), and killer whale ( Orcinus orca ) (Locke, 2008). Open tubules are associated with sensory ability in essay, mammals (Cuenin et al., 1991; Johnson and english Brännström, 1974; Panapoulos et al., 1983), though normally expressed only in pathologic conditions for most other mammals. Studies of nerve-associated tissue in water essays, the pulp of odontocete teeth are limited, though helpful in understanding function (Holland, 1994). Sensory innervation of mammalian teeth includes small myelinated A fibers and english charles lamb a majority of 1000 word on accountability unmyelinated C sensory fibers (Seltzer and Bender, 2002).
The A fibers are associated with dentin and english lamb the odontoblastic layer (Kimberly and Byers, 1988; Ikeda et al., 1997), while the C fibers are more uniform, though more densely populated in short answers, peripulpal areas and along blood vessels (Wakisaka et al., 1987; Hildebrand et al., 1995). Both A and C fibers contain the charles, neuropeptide calcitonin gene-related peptide, or CGRP (Silverman and Kruger, 1987; Fristad et al., 1994). In addition, C fibers contain the neuropeptide substance P (Casasco et al., 1990; Wakisaka, 1990). The presence of sensory-associated genes in the pulp is an essays, indicator for sensory function in mammalian teeth. Though there is a lack of english essays genomic information for the narwhal, techniques are available to measure the gene expression profiles using a universal array platform to want a wife essay, de novo sequencing that are informative to identify sensory-associated genes associated with the pulp (Velculescu et al., 1995; Unneberg, 2003; Roth et al., 2004). Other techniques using sequenced genomes of marine mammals similar to the narwhal as well as humans are also useful.
Evolutionary patterns of mammalian tooth anatomy and charles lamb morphology are driven by diet (Anapol and answers Lee, 1994; Jernvall et al. 1996; Teaford and Ungar, 2000; Evans et al., 2007; Thewissen et al., 2007; Lucas et al., 2008). Thus, comparing the foraging habits of male and female narwhals can potentially provide useful information about the erupted male tusk. Previous dietary studies comparing male and female narwhals from the Baffin Bay population have found no difference in stomach contents between the sexes (Finley and Gibb, 1982; Laidre and Heide-Jørgensen, 2005); however, smaller sample size of english female narwhals has limited these analyses, and stomach contents can be biased since they only provide information on kids, the most recent meal from a specific foraging area. Stable isotope and fatty acid analyses provide long-term integrated dietary information to investigate if dietary differences exist between male and female narwhals in the Baffin Bay population. Both techniques investigate chemical signals in english essays charles, animal tissues which have incorporated isotopic and narrative fatty acid values from their prey over various time frames, depending on the tissue. The nitrogen stable isotope (δ 15 N) provides information on an organism's trophic level, while the carbon stable isotope (δ 13 C) provides information on the animal's spatial foraging location, benthic versus pelagic or inshore versus offshore (Peterson and english Fry, 1987; Crawford et al., 2008; Newsome et al., 2010). Fatty acids are transferred relatively unmodified from prey tissues to predator tissues, and thus can also be used to determine and compare diet among groups of organisms (Iverson et al., 2004). Both analyses have been successfully used to investigate diet in marine mammals (Iverson et al., 2004; Newsome et al., 2010) including walruses ( Odobenus rosmarus ) (Dehn et al., 2007), bowhead ( Balaena mysticetus ), and gray ( Eschrichtius robustus ) whales, and why i want a wife essay the narwhal's closest relative, the beluga whale ( Delphinapterus leucas ) (Horstmann-Dehn et al., 2012). English Essays Charles Lamb? When the water kids, two analyses are used together and provide complementary results they can verify and confirm dietary interpretations, as was done in determining the primary prey for the beluga whale as arctic cod (Loseto et al., 2008, 2009) and for the bottlenose whale as Gonatus (Hooker et al., 2001).
Brännström's theory of dental sensitivity is the most widely accepted mammalian model to essays, explain this sensory ability and function for teeth. Changes to the interstitial fluid flow within dentinal tubules changes the conformation of odontoblastic cells connected to a pulpal nerve plexus which sends signals of sensory perception to the brain (Brännström, 1966). This theory explains the known ability of teeth to sense environmental stimuli, and water supports the evolutionary descriptions of essays charles tooth precursors as sensory organs (Lumsden, 1987). Human and other mammalian teeth are known to be capable of essay sensing external stimuli (Anderson, 1968; Haegerstam, 1976; Byers and Dong, 1983; Byers, 1984; Balam et al., 2005) such as temperature (Yamada et al., 1968; Jyväsjärvi and Kniffki, 1987; Ahn et al., 2012), pressure (Mengel et al., 1992), proprioception (Hassanali, 1997; Catania and Remple, 2002; Ozer et al., 2002), osmotic gradients, galvanic potential (Ramirez and Vanegas, 1989; Heyeraas et al., 1994), nocioception (Hu et al., 1978; Lisney, 1983; Narhi et al., 1984; Shigenaga et al., 1986; Byers et al., 1988; Iwata et al., 1998; Kawarada et al., 1999; Andrew and Matthews, 2000), and percussion (Ogawa et al., 2002; Watanabe et al., 2003). Sensory nerve fibers and essays lamb associated nerve bundles are also found in the pulp of other toothed animals (Pischinger and Stockinger, 1968; Weissengruber et al., 2005). This sensory ability serves many different functions, such as protecting teeth against environmental insults and water essays kids responding to age-related and essays charles lamb pathological conditions (Hildebrand et al., 1995). Because narwhals expose these open tubules during normal function, they are capable of sensing one or more of these variables. The erupted male narwhal tusk exhibits sensory ability based on physiologic response of changing heart rate associated with tusk exposure to alternating solutions of fresh and salt water, and the presence of sensory nerve-associated structures. A complete model of tusk sensory function, from why i want a wife, stimulus introduction to english essays charles lamb, brain perception, is the essay friendships, primary objective of this investigation. In vivo testing of this model to verify physiologic response to english essays, an introduced stimulus helps to confirm sensory ability.
Demonstration of this model includes: (1) an anatomical description of open channels in want essay, a porous cementum layer covering the erupted tusk; (2) further and more complete imaging of lamb a dentinal tubule network in various expressions of the narwhal tooth organ system, including the male erupted tusk, male and essay on regrets in life female unerupted tusks, and vestigial teeth in both sexes; (3) evidence of sensory-associated cells and tissue in the narwhal pulp and circumpulpal dentin; (4) detection of sensory-associated genes and antigens in the narwhal tusk pulp; (5) published findings of essays nerve connection at the base of the water essays kids, erupted tusk and continued connection to the maxillary division of the fifth cranial nerve to english, the brain; and (6) new results associating physiologic heart response in response to sensory stimuli of alternating fresh and high-salt water solutions introduced to on regrets, the tusk. An intact, erupted tusk (“A”) including pulp tissuetaken from charles, a male narwhal in 2003, 48 km from water, Pond Inlet, Baffin Island, in essays lamb, the eastern portion of Eclipse Sound, 24 km southwest of Cape Graham Moore, Bylot Island [72N 36′ 53.1″, 75W 37′ 56.4]was sawn in the field into 54 consecutive cross-sections measuring 3550 mm in axial length. Narrative? Overall erupted tusk length was 218.4 cm, with diameter increasing from 14.8 mm at the tip to essays lamb, 57.4 mm at essay, the base (Fig. 1A). Segments were fixed in 70% ethanol. Segment A-12 was processed to examine neuronal structures at the pulpal-dentin interface. The midpoint of segment A-12 was located 45.7 cm from the tusk tip, with a 2.78 cm mean outer diameter and english lamb 0.87 cm mean pulp diameter. Segment A-12 was secondarily fixed in 10% neutral buffered formalin, decalcified in EDTA, post-fixed in formalin, and paraffin embedded. Ten-micron-thick sections were mounted on SuperFrost slides for immunohistochemistry. (1A, top ): Sections A-12 and A-42 used in essay on regrets, SEM dentinal tubule analysis and charles surface imaging of short essay answers cementum, taken from an erupted male narwhal tusk; (1B, bottom ): Pulp sections used in english essays, immunohistochemistry analysis. An intact dental pulp was extracted from a narwhal tusk harvested near redundant text Kakiat Point, Arctic Bay, Baffin Island, in water essays, 2003 [72N 40′ 51.2″, 86W 41′ 16.9″].
The pulp was fixed in 70% ethanol for 2 months, then suspended in english lamb, 10% neutral buffered formalin at 4C under 20 g of tension to maintain extension. Overall dimensions of the essay development 1845, heavily vascularized dark red pulp were: length, 142.2 cm; diameters 0.3 cm (tip), 1.5 cm (mid), and 4.8 cm (base/apical tusk). The pulp base was bulbous and calcified. Ten-micron-thick paraffin sections were prepared from english charles, four separate segments of the pulp (C, 1.7 cm from pulp tip; F, 15.8 cm; I, 51.4 cm; and L, 90.8 cm). Segment L represented the format, pulp tissue at the approximate base of the erupted tusk that would be exposed to seawater (Fig. 1B). A fetus was also collected and head dissected into separate maxilla and mandible sections and stored in english essays, RNAlater ® solution (Life Technologies, Carlsbad, CA) and kept in ice until laboratory arrival. Tissue samples were brought under CITES Export Permits CA03FO1000005, CA05F01Q004, X0704-132/06, CA08F01Q0002, CA09F0W10001 and import permits 04US015415/9, 04US082589/9, 06US015415/9, US NMFS permits 932-1489-04 and 764-1703-00, and DFO transport permits A 02089, A 016505, 18469, 17378.
Examination of channels through a porous cementum layer and essay a dentinal tubule network patent through the full thickness of dentin was completed after sections were prepared. Samples for SEM were sectioned using a water-cooled diamond saw, and polished with a series of english essays lamb abrasives through 4000 grit with silicon carbide abrasive paper using a rotating metallography table. The surfaces were then cleaned to remove surface debris, using a mild HCl acid etch followed by rinsing in response, dilute sodium hypochlorite. The cleaned sections were then washed and english essays charles lamb dried in a vacuum desiccator and 1000 word essay and responsibility gold-sputter coated for essays charles SEM observation. Samples for cementum examination were prepared by taking cross-sections of friendships tusk segment A-42 sawn at 0.9-mm thickness with a precision low-speed water-cooled saw. Surface imaging of cementum was performed at english charles lamb, 50× objective magnification using a Keyence super resolution digital microscope model VHX-S90BE, equipped with a VH-Z20R/Z20W lens and water essays variable illumination adapter VH-K20. Reagents for essays charles lamb immunohistochemistry of doctrine 1845 neuronal markers included calcitonin gene-related peptide antibody (anti-CGRP, ab81887, abcam), 200kD neurofilament heavy antibody (ab17126, abcam), substance P antibody (#20064, Immunostar), and isolectin IB4-AlexaFluor 488 conjugate (Molecular Probes, Invitrogen). While specific for the indicated antigens in one or more mammalian species, none of these antibodies had previously been tested in narwhals, thereby precluding inference from negative findings. Sections were processed by microwave antigen retrieval for 15 min in citrate buffer solution, pH 6.0 (Antigen Unmasking Solution #H-3300 Vector Laboratories, Burlingame, CA).
Antibody localization involved peroxidase coupling and english essays charles lamb red color development with 3-amino-9-ethylcarbazole (ABC kit and ImmPACT AEC Peroxidase Substrate kit, #SK-4206, Vector Laboratories). Primary antibodies were diluted 1/50 to 1/1000 for incubation and were omitted for negative control sections; all sections were counterstained with aqueous hematoxylin. Slides were imaged by essays light microscopy with a Nikon Eclipse 80i at objective magnifications of 4× to 60× (oil). Since the narwhal has not been sequenced, a query of keywords “tooth” and “nerve” for the Tursiops truncatus genome was performed using the Ensembl database. English Charles Lamb? The genes NGFR, DLX2, TFAP2A, and GLI3were the only ones that resulted from the search. Primers were created for all four using the Roche Universal Primer Design software, though the QRT-PCR assay for GLI3 failed. Another approach to identifying additional sensory genes was based on the human genome: FAM134B and WNK1 (formerly HSN2) genes have been shown to be involved in hereditary sensory neuropathy type II, which has a pathology for response the reduction or loss of the sensory perception of pain, temperature and touch (Kurth, 2010). These genes were cross-referenced to the Tursiops truncatus genome, and to our surprise, based on the Ensembl database search, there was a sequence similarity for these two genes.
The narwhal pulp tissue was stored in DNAgard (Biomatrica, San Diego, CA) and the control samples (muscle tissue around the blow-hole and gingiva from the inner portion of the mandible) were placed in RNAlater solution following the charles lamb, respective company protocol within 15 min of essay christian doctrine access to a fetus collected at english charles lamb, Kakiat Point, then all samples were stored at −20C until arrival at HC-LITT, where they were stored at −80C. Sections of these pulp and control samples were obtained from multiple regions and weighed prior to development christian, homogenization. The Covaris t-PREP™ and S220 Focused-ultrasonicator (Covaris, Woburn, MA) were used to homogenize the pulp and control samples. Prior to essays lamb, Adaptive Focused Acoustics™ on the S220 instrument, 250 μL of short essay format answers Qiagen Buffer RLT (Qiagen, Valencia, CA) was added to the samples. After processing, 750 μL of TRIzol LS was added to extract the RNA from the essays charles lamb, samples using the standard three-step phenol/chloroform extraction method according to the manufacturer's instructions and prior to a wife response essay, making cDNA. cDNA was synthesized from 1.5 µg of charles lamb total RNA for each sample using Roche reverse transcriptase. All assays were prepared using standard conditions in a master mix solution without any effort at assay optimization. For each data point, there were three replicates. The sensory and essay development doctrine housekeeping genes were designed based on the Tursiops truncatus and Stenella coeruleoalba genomes, respectively.
The Ensembl database (http://uswest.ensembl.org) was utilized to project transcripts involved in sensory nerve functions by english essays charles using the short answers, closest relative of the english essays, narwhal whose genome had been sequenced ( Tursiops truncatus http://uswest.ensembl.org/Tursiops_truncatus/Info/Index [Database version: 71.1]) (Hubbard et al., 2002; Flicek et al., 2012). The gene accession numbers were used in format, designing the primers for the housekeeping genes (Spinsanti et al., 2006). The Roche Universal ProbeLibrary probes and target-specific PCR primers were selected using the ProbeFinder Assay Design software (https://qpcr.probefinder.com/organism.jsp, “Other organism” selection) (Rozen and Skaletsky, 2002; Kuo et al., 2006). Please refer to Table 1 for primer sequences. The primers were used to confirm relative changes in mRNA levels by lamb QRT-PCR using the why i want essay, Roche 480 LightCycler. Reactions were performed in microliter reaction volumes for six genes, using 1 μL of cDNA under the following conditions: 95C for 10 min, followed by essays 45 cycles of 95C for 10 s and essays 60C for 30 s (Kuo et al., 2006). Expression data for the sensory genes were normalized relative to the housekeeping genes (Murphy et al., 2003; Khimani et al., 2005). Raw data were obtained and analyzed using the SBI software (System BioSciences, Mountain View, CA). Charles? The comparative C T method (ΔΔ C t) was chosen to further calculate the short format, relative expressions between the different groups. Carbon (δ 13 C) and nitrogen (δ 15 N) stable isotope analyses were performed on male ( n = 91) and female ( n = 91) narwhal skin samples collected by Inuit hunters in the Canadian High Arctic from 1982 to 2009. Narwhal skin subsections of essays charles lamb approximately 0.5 g were freeze-dried, lipid-extracted, and analyzed at the University of Windsor for stable isotopes.
Burning of fossil fuels emits anthropogenic CO 2 that is depleted in 13 C, resulting in increasingly depleted C animal tissue signals over essay christian doctrine the industrial period: this pattern is known as the essays, Suess effect (Sonnerup et al., 1999). Carbon values were corrected to account for this depletion by calculating the atmospheric carbon value ( C atm ) in 1982 (the oldest sample included in the analysis) using δ 13 C atm = −6.429 to 0.0060 exp (0.0217[ t −1740]), where t is the year (Feng, 1998). Atmospheric δ 13 C atm was calculated for each year samples were collected, and format answers the difference between that value and the 1982 δ 13 C atm was subtracted from the measured δ 13 C to obtain a corrected carbon value. Lamb? For the analysis of 1000 fatty acids, blubber samples from female ( n = 56) and male ( n = 56) narwhals harvested near the english, community of essay doctrine 1845 Pond Inlet, Nunavut, Canada were used. English Essays Charles Lamb? Lipids were extracted from narwhal blubber samples for fatty acid analysis using the why i response, modified Folch procedure (Folch et al., 1957; Budge et al., 2006) and converted to fatty acid methyl esters (FAME). FAME were analyzed using the Agilent Technologies 7890A GC system. T-tests were used to english, test for differences in δ 13 C and δ 15 N between male and female narwhals.
Fatty acid values were converted to percentages, divided by water the mean of the sample, and log-transformed (Kenkel, 2006). A principal component analysis, which included 33 dietary fatty acids (Iverson et al., 2004), was followed by english charles a multivariate ANOVA on the first five principal components (cumulatively describing over 70% of the data) to determine if males and narrative friendships females differed significantly in their fatty acid values. English Essays Lamb? All statistical analyses were conducted using JMP 7.0.1. Alternating high-salt (41 ppt) and fresh-water solutions were used as stimuli to test the physiologic response of the narwhal tusk to environmental sensory stimuli. Selection of the stimulus was based on the potential relevance and importance of salt ions to why i want a wife response essay, the narwhal's ocean environment. A tusk jacket was constructed from a 10.2 cm diameter by 45.0 cm length clear acrylic tube with a 0.64 cm wall thickness, which was cut in english essays charles, half longitudinally and hinged on one seam to facilitate immediate closure at the other seam.
At each end a 5.7 cm wide by essay on regrets in life 3.8 cm thick piece of open-celled foam was glued to the inner surface of the tube, with an english essays charles, overall diameter of and responsibility 10.2 cm and an inner open diameter of 2.8 cm, to form a centering gasket around the tusk at charles, the tube ends. A one-way hose valve was fastened to the center of the tusk jacket, and Versilic Formulation SPX-70 I.B. tubing connected the jacket to water, an MSR 6-L Dromedary solution storage bag. Quick-connect valves allowed easy exchange of solution storage bags filled with fresh-water or high-salt solutions. The Midmark Holter IQ electrocardiograph (ECG) with a five-lead montage and eight-second data averaging was used for the six male narwhal samples tested in Tremblay Sound (TS) near the base of the Alpha River outside Pond Inlet [72N 21′ 38″, 81W 05′ 59″], two each in english essays, years 2010, 2011, and 2012. Heart rate response was selected as a physiologic indicator, since sudden increases in salt ion concentrations would presumably be interpreted by narwhals as a sign that ice entrapment was threatening. Thus, during the introduction of short essay high-salt solutions narwhal heart rate was expected to rise due to essays charles lamb, physiologic response to rapid ice formation; conversely, heart rate was expected to fall during the introduction of fresh water, as this stimulus is associated with the waters of essay development christian summer inlets where ice is melting and there are fresh-water run-offs. The narwhals were captured from land-based camps using an anchored net set perpendicular to the shore, and english essays lamb after capture brought to shore using standard techniques established for tagging (Dietz et al., 2001, 2008). Why I Want A Wife Response? After the english lamb, narwhals were stabilized, subdermal electrodes were inserted in a five-lead montage. Electrode leads for ECG data collection were Grass Technologies subdermal Model F-E7 right-angled, 10 mm, 30-gage monopolar needles with 244 cm lead wires. IQ Manager software with real-time signal collection was used with the essay, Holter IQ. Eight-second interval averages were used in the final analysis.
Solutions of 41 ppt salt-water were made by english essays charles lamb adding Instant Ocean™ saltwater synthetic salt crystals to narrative friendships, inlet brackish sea water (1822 ppt); salt concentration was confirmed using the Omega CDH-287 meter. Fresh-water solutions were gathered from nearby glacial run-off streams that were used as drinking water. McCormick red and green food coloring dyes were added (10 drops/gallon) to identify the english essays charles, two different solutions, and to verify the why i a wife essay, clearing of one solution in the tusk jacket and displacement by the alternating solution. Essays? The two dye colors were interchanged between high-salt and fresh-water solutions to why i want a wife essay, eliminate the possibility of lamb food coloring sensitivity as another variable detected by the tusk, or color dispersion observed by the animal. The ECG was turned on after electrode attachment. After a few minutes of heart monitoring, the tusk jacket was attached at or near the base of the tusk, and 1000 essay alternating solutions of high-salt and fresh water were injected into the tusk jacket. Temperature of these stored solutions was kept at ambient sea water level by essays charles keeping them in Dromedary™ bags submerged in nearby seawater. Time periods for the solution injections were varied between animals to essays kids, account for any effect on english essays lamb, changes in heart rate response linked to capture time. Essay 1845? Heart rate was monitored continuously during the solution exchange and english essays charles dwell times until all experiments were completed. The Midmark Holter IQ™ instrument allowed real-time heart rate data to be stored and later segmented into 8-s averaging for the sample of six male narwhals.
The data recording was electronically marked at the beginning of each stimulus period to provide an accurate association of 1000 word essay recorded heart rate to stimulus. English Essays Lamb? The five-lead montage provided a manageable configuration of electrodes for field use, and 1000 and responsibility very few interruptions in english essays lamb, data were experienced due to essay development christian doctrine 1845, displacement of the subdermal electrodes. An ideal heart beat wave form was identified to correlate with data recording and analysis (Fig. English Essays? 2). Animal health and safety is narrative, always the primary concern, and all experiments cease and essays the animal is released at the first sign of overt stress when indicated by the monitoring veterinarian. Fortunately, none of the essay, animals exhibited problems requiring early release, and most experiments, tagging, and lamb specimen collections were completed within 30 min.
Experiments and research were conducted under Canadian Department of 1000 Fisheries and Oceans' Animal Care Committee permits FWI-ACC-20082009-007 and FWI-ACC-2008-2009-008 and Animal Use Protocol permits FWI-ACC-2009-024, FWI-ACC-2010-008, FWI-ACC-2011-016, and FWI-ACC-2012-009. Essays Charles? Research protocol was established in the United States under the Harvard Medical Area Standing Committee on Animals, Animal Experimentation Protocol 04384. The Hunters and Trappers Organization (HTO) in Pond Inlet, Nunavut voted to christian, approve these research protocols in TS. A generalized least squares split-plot nested analysis was chosen to account for the unbalanced experimental design where there was variation in the length of the stimulus periods, and english essays there was a reasonable likelihood that residuals would correlate between stimuli within each individual animal. Statistical analysis on the six animals captured at water essays kids, TS was done with whales and water treatments as the primary fixed factors and split-plots nested within whales and water treatments, to determine parameter estimates and the significance levels of the fixed factors and charles factor interactions, fitting the model: µ + W i + T j + (WT) ij k + ε ijkl + γ (ij). This model included both split-plot and random error to essay development christian 1845, account for the unbalanced experimental design where treatment intervals were not equal in length or number.
The model also accounted for the assumption that residuals would be correlated within each individual animal. No attempt was made to identify or exclude outliers in the data, although a few data points appeared to charles lamb, be potential outliers. Tukey Honest Significant Difference (HSD) simultaneous confidence intervals were generated at 95% to evaluate the sizes of the differences among the whales' mean heart rates. A Student's t 95% confidence interval estimate was constructed to evaluate the size of the difference between treatment mean heart rates. 1000 On Accountability And Responsibility? Statistical analyses were conducted using JMP Pro 10.0.0. Results confirmed multiple indicators that support the hypothesis of a sensory pathway in the erupted narwhal tusk. Scanning electron micrographs of patent dentinal tubule networks provide evidence of the pathway for sensory conduction in multiple forms of essays narwhal tooth expression, including both male and friendships female erupted tusks, unerupted tusks, and vestigial teeth (Fig. 3). Narwhal electrocardiogram wave components from essays charles lamb, TS sample Whale 3. P wave = atrial depolarization. Short Essay Answers? PR interval = time interval between onset of atrial depolarization and essays charles lamb onset of ventricular depolarization. Essay Friendships? QRS complex = depolarization of ventricles, consisting of the Q, R, and english lamb S waves.
QT interval = time interval between onset of ventricular depolarization and end of ventricular repolarization. RR interval = time interval between 2 QRS complexes. T wave = ventricular repolarization. ST segment plus = ventricular myocardial depolarization. U wave = relaxation of ventricles.
Organized and open channels were observed in tusk cross-sections, providing direct communication between the outer cementum surface of the tusk and the dentin-cementum interface (Fig. 4). 1000? The average channel diameter is 31 ± 14 (S.D.) μm, and the open areas comprise about 0.5% of the english lamb, cementum. This sinusoidal channel network appears to surround bundles of short answers Sharpey's fibers, and represents a possible pathway for sea water conduction through the cementum layer to the underlying dentin. Scanning electron micrographs at 1000× showing dentinal tubule orifices for english lamb ( A ) male unerupted tusk and ( B ) female erupted tusk and at want response essay, 500× showing polished dentin sections of ( C ) adult male tusk (inner third polished and essays lamb etched), ( D ) adult female tusk and ( E ) unerupted tusk. In Life? (Orientation of micrographs B-D, pulpal surface to the left and outer surface to the right). Scale bars = 10 microns.
Putative nerve fibers in narwhal pulp and dentin were identified by charles immunohistochemical detection of word neuronal markers. CGRP and substance P are neuropeptides expressed by neurons in dental pulp with important roles in english essays charles lamb, physiology and pathology (Caviedes-Bucheli et al., 2006). CGRP+ fibers were found localized to peripheral pulp (Fig. 5), and while few were surrounded by water essays connective tissue or adjacent to vessels, most were typically within 100 μm of the pulp margin. Essays Charles Lamb? This outermost pulpal region borders the odontoblast layer. Antibody to 200 kD neurofilament heavy chain, another neuronal marker reported in human dental pulp (Henry et al., 2012), failed to essay on regrets, localize to any structures in narwhal pulp (not shown). Organized channels traverse the full thickness of narwhal cementum. Scale bars = 500 microns. Left panel: numerous channels (white arrows) averaging 31 ± 14 µm in diameter are observed throughout the cementum layer in this cross-section of tusk specimen A-42 (see Fig. 1). The sinusoidal channel shape results in a segmented appearance in essays lamb, the image plane.
The outer surface of the tusk (black arrow) is 6.3 mm from the cementum-dentin boundary at the right margin of this panel. Right panel: rotated image showing the cementum-dentin boundary where the fluid-filled dentinal tubules are in close proximity to the channels. Sharpey's fibers (S) that anchored the erupting tusk to alveolar bone during development appear as dark, hypomineralized collagen bundles. Substance P was clearly present in the peripheral pulp (Fig. Word? 5). We observed extensive tracts of essays charles red substance P+ granules with diameters of 0.52 μm (Fig. 6, panels A, B, D). Want Response Essay? Substance P+ granules were found in english essays, four pulp segments representing the major linear dimensions of the pulp from water kids, tip to base of the erupted tusk (C, 1.7 cm from pulp tip; F, 15.8 cm; I, 51.4 cm; and L, 90.8 cm). In all pulp segments (where cross-sectional dimensions ranged from 0.4 to 15 mm), the substance P+ neurons were largely restricted to the outermost 200 μm of english essays circumpulpal tissue in closest proximity to the odontoblast cell bodies. Consistent staining of small discrete loci always at the pulp periphery and in the exact same positions through consecutive serial sections help to confirm the accuracy and essay precision of the technique. The granule distribution and size is essays charles lamb, suggestive of parallel, oriented dendrites or peripheral processes of sensory neurons, which are similar to the boutons en passant (beads on a string) morphology of substance P+ neurons in essays, the CNS (Behan et al., 1993).
The substance P+ granules provide strong evidence of peripheral innervation with potential for nociceptive transmission of excitatory impulses to the CNS. Other roles could involve substance P-mediated regulation of pulpal blood flow and extravasation and other responses to essays lamb, inflammation. Substance P, a peptide product of the Tachykinin 1/TAC1 gene, acts on christian doctrine, its receptor NK1R (Tachykinin receptor 1/TACR1) expressed by english essays charles lamb neurons and endothelial cells. Substance P released from pulpal sensory nerves during experimental tooth pain is implicated in nociceptive pain transmission (Avellán et al., 2008) as well as inflammatory responses to pain, including increased pulpal blood flow (Kim, 1990). CGRP immunostaining (red color) of putative nerve fibers (white arrows) in peripheral zone of narwhal pulp adjacent to circumpulpal dentin. Panel APulp Segment F; panel BSegment L, upper left arrow is margin of large artery with smooth muscle wall; panel CSegment F; panel DSegment L. Control sections with primary antibody omitted showed no red reaction product. Water Essays? Scale bars = 20 microns. We examined substance P expression in essays charles lamb, the layers of odontoblasts, predentin, and dentin immediately adjacent to the peripheral pulp tissue. Odontoblasts were not evident at the pulp surface (Figs. 4A and essays kids 6B) because they remained attached to the circumpulpal dentin when the intact pulp was forcibly withdrawn from the tusk.
Sections from Segment A-12 of a tusk containing its pulp faithfully preserved this interface (Fig. Essays Charles Lamb? 6DF). There is evidence of substance P+ nerve fiber adjacent to odontoblast cell bodies, and further evidence of substance P+ fibers in close proximity to odontoblastic processes in the zone of dentinal tubules (Fig. On Regrets? 6EF). This anatomical association supports the charles lamb, possibility of odontoblast-initiated sensory transmission in the narwhal tusk. Substance P immunostaining (red color) of putative nerve fibers in narwhal pulp and dentin. Panel A, pulp segment L showing abundant red SP+ granules at kids, pulp periphery (arrow); panel B, segment I showing linear arrays of SP+ granules near pulp periphery (arrows); panel C, segment C showing small cluster (arrow) of red SP+ granules at pulp periphery; panel D, segment I showing SP+ nerve fiber (arrow) adjacent to large artery with smooth muscle wall (sm); panel E, tusk segment A12 showing SP+ nerve (n) in essays charles lamb, peripheral pulp adjacent to odontoblast cell bodies (o) and word essay on accountability and responsibility dentin (d) with characteristic dentinal tubules (t); panel F, tusk segment A12 showing linear arrays of essays lamb SP+ granules (arrows) in odontoblast (o) layer of dentin approximately 200 microns from the peripheral pulp. Scale bars = 100 microns (A, B, D, E) and 10 microns (C, F). Top : Fold-change of sensory genes when compared to muscle tissue. Bottom: Fold-change of sensory genes when compared to narrative essay friendships, mandibular soft tissue.
All the charles, genes were normalized using the housekeeping genes (GAPDH, RPL4, RPS18, and YWHAZ). The gene expression profiles for sensory genes were calculated based on fold-change. DLX2, FAM134B, NGFR, and TFAP2A genes were highly expressed in the pulpal tissue when compared to both muscle and mandibular soft tissues (Fig. 7). When compared with pulpal tissue, the fold-change ranged from 1.64 (for FAM134B) to 21.03 (for DLX2), and narrative three of the genes that had a fold-change greater than 16.5 when compared to essays lamb, muscle were highly expressed. Since the mandibular soft tissue might contain sensory nerves the gene expression profiles were not as significant, and these genes had fold-changes greater than 2.5. The gene expression profile of WNK1 was marginally higher in pulpal tissue compared to muscle tissue, and slightly down-regulated when compared to mandibular soft tissue (Hubbard et al., 2002; Kurth, 2010; Flicek et al., 2012). On the other hand, NGF was the only gene that was down-regulated compared with both muscle and mandibular soft tissue; thus, the word essay and responsibility, expression was higher in both muscle and mandibular soft tissue than in the pulp. Carbon and nitrogen stable isotope values and dietary fatty acid percentages are both biomarkers that provide an english essays, indication of diet. Biomarkers were significantly different between male and female narwhals from the Baffin Bay (High Arctic) population, suggesting the why i want, sexes may have different foraging habits.
Carbon and nitrogen stable isotope data were normally distributed, as described by essays charles lamb normal quantile plots, and christian doctrine variances were homogenous (Levene's test: Carbon, F 1, 432 = 0.433, P = 0. 51; Nitrogen, F 1, 432 = 0.28, P = 0.60). Male narwhals ( n = 278) had significantly higher δ 13 C ( t -test: t = 4.82, P 0.0001) and δ 15 N ( t -test: t = 3.5, P = 0.0004) values compared to female narwhals ( n = 156). A multivariate ANOVA on the first five fatty acid principal components found there was also a significant difference between male and female narwhals (MANOVA: F 5, 106 = 2.42, P = 0.041, n = 56). This significant difference between stable isotope and fatty acid values for male and female narwhals supported divergent sexual foraging habits.
A plot of the actual versus predicted heart rates indicated that there may have been a few observations that were outliers, but these observations were not eliminated from the data set of six male narwhals from TS (Table 2) since they had little impact on the overall analysis. Essays Charles Lamb? A summary of fit of the actual and essay format answers predicted heart rates resulted in an R 2 value of 0.87. A test of the fixed effects indicated that the difference among the whales was statistically significant ( P 0.0001) and charles that the treatment difference (high-salt versus fresh-water exposure) was significant ( P = 0.0008); however, there was no significant two-way interaction between the whales and treatment ( P = 0.36). Narrative Friendships? A plot of the residuals showed a random pattern around zero, indicating that the assumption of equal variances was likely met. The Tukey HSD set of english lamb confidence intervals of the differences (Table 3, bottom) indicated that there was no difference in the estimated mean heart rates between whales 1 and 2, whales 1 and 3, and essay format whales 2, 5 and english charles lamb 6 at the 95% confidence level. Whale 4 was significantly different from all others. The least squares means differences in 1000 word on accountability and responsibility, Student's t test of the salt and essays charles fresh-water treatment conditions (Table 3, top) resulted in an overall mean estimated heart rate of 60.42 bpm for the 41 ppt salt-water stimulus and an estimated 52.56 bpm for the fresh-water stimulus (Table 4), and these were significantly different ( P = 0.0008). Doctrine? The estimated difference between the essays lamb, two means, with 95% confidence, was 7.86 bpm. The least squares means analysis of the whale-treatment interaction had a P -value of 0.36, indicating that all such confidence intervals of mean differences would overlap. These results demonstrated a significant and repeatable difference in heart rate, with higher heart rate being associated with the 41 ppt salt-water exposure and lower heart rate with the fresh-water exposure (Fig. 8).
This change in heart rate occurred in all six animals, though there were differences in doctrine 1845, the heart rates measured between the charles lamb, different animals. The magnitude of change for the salt stimulus mean was between 3.7 and 12.0 beats per minute. Mean heart rate measured for narrative essay each of the six male narwhals while the tusk is stimulated with either 41 ppt salt water or fresh water. Sensory model of the erupted male narwhal tusk showing, from bottom left, the essays charles lamb, introduction of narrative essay friendships water gradients penetrating cementum channels connected to patent dentinal tubules through the full thickness of the dentinal layer, connecting to odontoblastic processes and cells at the base of the tubules and at the periphery of the pulp, which stimulate nerve tissue connecting the base of tusk tissue to the maxillary branch of the charles lamb, fifth cranial nerve to the brain. Also pictured at the bottom right are pulp peripheral nerve-associated substance P and CGRP. Human and other mammalian teeth have sensory function and exhibit a highly adapted ability to why i essay, sense variables including temperature, ion concentration gradients, physical pressure during loading, and proprioceptive position. It is therefore reasonable to expect that this function of dentition is also found in english essays charles lamb, the narwhal.
Anatomic form and physiologic field results are consistent with Brännström's hydrodynamic theory of teeth, the most widely accepted theory of sensory and pain perception associated with exposed dentinal tubules (Brännström, 1966, 1967). Essay Friendships? The theory hypothesizes that interstitial fluid within these tubules can be influenced by variables of temperature, pressure, hydration status, and electrochemical and charles osmotic gradients. 1000 Word And Responsibility? Solutions introduced at essays charles lamb, the tusk surface have the effect of narrative essay penetrating a porous cementum outer layer through a series of open and organized channels that create fluid flow within the dentinal tubules, outward during the exposure to charles, high-salt solutions and inward after exposure to a wife, fresh water. Mechanical effects on odontoblastic processes within these tubules, and close associations of odontoblasts with nerve endings in the outer layer of pulpal tissue (Fig. 9), are involved in charles lamb, the initiation of sensory perception.
Direct exposure to the environment of the patent dentinal tubule network in kids, narwhals is unusual. In contrast, the beluga ( Delphinapterus leucas )the closest relative to lamb, the narwhalexhibits the 1000 word on accountability and responsibility, more characteristic occluded dentinal tubules within the essays, dentin layer, preventing any fluid conduction from the essay, enamel-covered tooth surface to the ocean environment (Sergeant, 1959 and 1973). Primer sequences used in lamb, QRT-PCR experiments lists the Roche Universal Probe number and 1000 word essay corresponding forward, reverse and amplicon sequences for english essays charles the Universal Probe Library reactions. The sensory and housekeeping genes were designed using the Tursiops truncatus and Stenella coeruleoalba genomes, respectively. Tubules are present in all mammalian dentin and serve to narrative essay friendships, supply the english lamb, physiologic fluids that maintain the integrity of collagen that forms the collagen fibril network within dentin (Pashley, 1996; Kinney et al., 2003). It is well known that in human root-canal-treated teeth, where the pulpal tissue has been removed and thus the essay, internal supply of physiologic fluid, the english charles lamb, dentin becomes more brittle and loses elasticity over time (Jameson et al., 1993; Papa et al., 1994). This same physiologic function of short essay answers maintaining collagen toughness within narwhal dentin would be even more critical at english essays charles lamb, the high bending stresses experienced within a tusk. Tensile stresses are greatest near the base and on on accountability, the surface of a cantilevered beam, such as a narwhal tusk. Tissues with very high fracture toughness, such as the highly fibrous and english essays charles collagenous dentin of the tusk, are best designed to respond to why i want response, these stresses without fracturing. This tubular tusk structure of toughened dentin is a wonderful example of evolutionary engineering, where environmental demands have resulted in optimized organ design.
This explains the high density of english charles tubules observed and essay development christian doctrine the fact that narwhal tusks maintain a pulpal vascular network for nearly their entire length. The unique feature of narwhal tusk dentin, however, is the frequent and essays normal communication of water essays these tubules with the exposed surface of the tusk. In other mammalian teeth, surface layers of charles enamel or cementum occlude tubules, and why i occluded tubules frequently occur within the english charles, dentin layer, so that exposure of tubules only occurs as a result of damage or pathology. Want A Wife Response Essay? The narwhal tusk has no surface enamel layer (Ishiyama, 1987), and charles the cementum observed on the surface is highly porous with a network of open and organized channels surrounding abundant Sharpey's fiber bundles of hypomineralized collagen, thus providing the essay in life, patent communication needed to sense surface stimuli. Dissection and imaging of male and female narwhal heads reveal connection of the english, maxillary division of the fifth cranial nerve from the brain to the tusk. Water Essays Kids? This connection is supported by intracranial dissections, tusk alveolar examination, and visible pathways of nerve conduction observed by dissection and charles lamb radiographic imaging (Tyler, 2006; Nweeia et al., 2009). Narwhal pulp has been reported to have features similar to other mammals (Dow and short essay Hollenberg, 1977).
Nerve-associated tissue discovered in narwhal pulpal tissue (Figs. 4 and english charles lamb 5) also provides evidence that the Brännström hydrodynamic theory of sensory function could apply to the narwhal tusk. Transmission of dental sensory or nociceptive (pain) signals to the CNS involves Aδ and C-type fibers. Rapid, sharp pain is associated with Aδ fibers, while dull, radiating pain typically involves C-type fibers, depending on format, the type of stimulus. The Aδ fibers account for the sensitivity of dentin, whereas C-type fibers are activated only if external stimuli reach the pulp proper (Narhi et al., 1992).
Hydrodynamic sensory perception in the narwhal tusk is supported by our findings of charles lamb extensive innervation and a wife expression of substance P, a known nociceptive neurotransmitter, in the peripheral region of the pulp. The close apposition of odontoblasts and substance P+ fibers is consistent with odontoblast-dependent sensory transmission initiated by lamb fluid flow in dentinal tubules. Essay Format Answers? Recent reports of mechano-sensitive ion channels such as TRPV14 and essays charles TREK-1 in odontoblast membranes, and essay format PC1 and PC2 associated with the odontoblast primary cilia (Magloire et al., 2009, 2010), provide possible additional mechanisms for odontoblast-mediated sensory transmission. Significant presence and detection of a set of sensory-related genes in the pulp (including DLX2, FAM134B, NGFR, TFAP2A, and NGF), either highly expressed or less expressed relative to english lamb, control samples of muscle and mandibular soft tissue, suggest a genetic basis for sensory function. The results of stable isotope and essay on regrets in life fatty acid analyses suggest sexual foraging differences that may relate to additional functional attributes of the essays, tusk. Narrative Essay? Stable isotope values in large mammal tissues can reflect the lamb, previous year's diet (Sponheimer et al., 2006), whereas fatty acids in blubber have turnover rates on the order of 1.53 months (Nordstrom et al., 2008). The majority of our samples were collected in why i want essay, July and August.
Thus, fatty acid signatures would reflect dietary habits during the late spring to early summer period, which may be after the breeding season in either mid-April (Best and essays lamb Fisher, 1974) or May-June (Heide-Jørgensen and Garde, 2011). Fatty acid values varied between males and short essay females, although much less strongly than stable isotopes, which may suggest males and females are foraging more similarly during the mating season or that both sexes limit feeding at this time. While more evidence is certainly needed, there remains the possibility that tusk sensory function could play a role in locating food sources. Physiologic data from six live-captured male narwhals suggest statistically significant sensory perception to high-salt and essays lamb fresh-water solutions (Fig. 7). It is interesting to water kids, note that Whale 1 presented with one-third of its tusk broken off. Results indicate this tusk damage compromised the ability to distinguish between high-salt and freshwater solutions. Whale 2 also presented with a broken tusk, but only a minor 23 cm loss of length, and thus sensation of salinity was only slightly compromised when compared to Whales 36. An established sensory pathway has thus been discovered from the external ocean environment through porous cementum, penetrating patent tubules conducting fluid shifts to sensory cells in the pulp, and transfer to essays charles lamb, the maxillary division of the fifth cranial nerve which connects to the base of the brain.
This anatomic model (Fig. 9), combined with field experiments showing sensitivity to high salinity gradients and associated increases in physiological heart rate response, show that the narwhal tusk organ has sensory ability. This is an water kids, organ with the ability to sense characteristics of its external environment. Essays Charles Lamb? More common associations of tooth sensitivity to their environment are linked to 1000 essay, protection. Teeth sense stimuli that signal a threat to their function or integrity and essays lamb the animal responds in a normal “fight or flight” reaction to avoid or neutralize the want a wife essay, stimulus. Such stimuli are often episodic, appearing when the threat is present and not as a normal part of the essays charles, tooth's environment. The narwhal is the only example documented where teeth are shown to have the ability to constantly sense environmental stimuli that would not necessarily be considered a threat.
Unusual tusk microanatomy provides a unique means of assessing the essay christian doctrine 1845, whale's environment by what appears to be their normal and commonly employed mechanism. The dentinal tubule network in the embedded (unerupted) tusk antemere opens the possibility that it may serve as a “reference tusk,” bathed in physiological fluid and used as a set point for sensing “high” versus “low” salinity by lamb the erupted “sensory tusk.” Tested sensitivities to high-salt and fresh-water solutions suggest that these dentinal tubules may also sense changes in temperature, pressure, hydration status, and/or electrochemical and osmotic gradients. Further examination may likely reveal other variables that can be sensed and interpreted, according to Brännström's hydrodynamic theory, by this extraordinary tooth organ system. Future experiments that test additional stimuli would provide a better understanding of the broader capability of this sensory organ system. The current evidence suggests that the essays, tusk may have a multitude of specific and perhaps interacting functions. Its microanatomy and sensory ability compellingly point to function as a sensory organ system.
Although erupted tusks may present in both sexes, they are mostly found in males and thus are more likely associated with sexual selection. Essays Lamb? The microanatomy of the female tusk reveals a similar dentinal tubule network. However, their gross morphology of a more dense proportion of hard tissue with a very limited pulpal canal suggests that they may not have the same sensory ability. Other gross morphological differences in erupted female tusks, which are present in approximately 15% of the essay format, population (Roberge and Dunn, 1990), have also been documented (Clark, 1871; Pederson, 1931). Essays? No field experiments were conducted on kids, females with tusks, as none were part of the essays, sample caught during this investigation. The use of teeth in why i want a wife response essay, intrasexual selection or “male-to-male rivalry” and competition amongst odontocetes has been documented in the literature, and may have relevance to narwhal tusk function.
New evidence of english essays lamb sensory ability suggests that intersexual selection or “mate choice” should be considered as a mechanism for sexual selection. Several hypotheses can be suggested for “mate choice” based on the sensory ability of the erupted male tusk. The first is that the tusk might detect waters where females in estrus may be gathered or attracted. A second is that the tusk could also detect waters where females are foraging, as observed from fatty acid and stable isotope results. As an expression of fitness, a third hypothesis is that some males may be better able to find sources of foods needed for newborn calves. Essay Christian Doctrine? The results of stable isotope and fatty acid analyses suggest sexual foraging differences that may relate to additional functional attributes of the tusk. Tusk sensory ability as a retained function from an evolutionary precursor is a fourth hypothesis; a fifth is that the tusk is a currently evolving sensory organ. Differences in english essays charles, tooth position and morphology suggest other functions such as species recognition, as they do in in life, other odontocetes (MacLeod, 2000). Since behaviors linked to mating are difficult to collect throughout the year, and a significant time period from english essays charles lamb, late October to March can be forbidding due to kids, weather and ocean conditions including moving pack ice within Baffin Bay in the early spring (Mansfield et al., 1975), observations of narwhal behavior will continue to be difficult to gather.
Such observations may be essential, however, for a more complete understanding of the english essays charles lamb, significance of tusk sensory function. Thanks are extended to the National Science Foundation Offices of Polar Programs and Integrative and Organismal Systems, elders and hunters from the High Arctic communities of Nunavut, Canada and northwestern Greenland, and short essay format the Arctic field teams and The National Geographic Society. calcitonin gene-related peptide. fatty acid methyl esters. honest significant difference. hunters and trappers organization. scanning electron microscopy. Copyright 2014 Wiley Periodicals, Inc. narwhal; tusk; Monodon monoceros ; tusk functional significance; Brännström hydrodynamic theory; tooth gene expression; pulp tissue; tusk histology. Issue online: 18 March 2014 Version of charles lamb record online: 18 March 2014 Manuscript Accepted: 15 January 2014 Manuscript Received: 8 November 2013. Ahn DK , Doutova EA , McNaughton K , Light AR , Narhi M , Maixner W . 2012 . Functional properties of tooth pulp neurons responding to thermal stimulation . J Dent Res 91 : 401 406 . CrossRef | CAS | Web of Science® Times Cited: 2 Anapol F , S. Lee . 1994 . Essay? Morphological adaptation to diet in platyrrhine primates . Am J Phys Anthropol 94 : 239 261 . Charles Lamb? Wiley Online Library | PubMed | CAS | Web of Science® Times Cited: 62 Anderson DJ . 1968 . Studies on sensory mechanisms in teeth . Narrative Friendships? Ann R Coll Surg Engl 43 : 331 340 . PubMed | CAS Andrew D , Matthews B . 2000 . Displacement of the charles lamb, contents of dentinal tubules and sensory transduction in intradental nerves of the cat . J Physiol 529 ( Pt 3 ): 791 802 . Wiley Online Library | CAS | Web of Science® Times Cited: 33 Avellán NL , Sorsa T , Tervahartiala T , Forster C , Kemppainen P . 2008 . Experimental tooth pain elevates substance P and matrix metalloproteinase-8 levels in human gingival crevice fluid . 1000 Essay On Accountability? Acta Odontol Scad 66 : 18 22 . CrossRef | PubMed | CAS | Web of Science® Times Cited: 4 Balam TA , Yamashiro T , Zheng L , Murshid Ahmed S , Fujiyoshi Y , Takano-Yamamoto T . 2005 . Experimental tooth movement upregulates preproenkephalin mRNA in the rat trigeminal nucleus caudalis and oralis . Brain Res 1036 : 196 201 . CrossRef | PubMed | CAS | Web of Science® Times Cited: 8 Beddard FE . 1900 . A book of whales . New York : Putnam and Sons.
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Washington, D.C .: Smithsonian Institution Scholarly Press. Essays Charles Lamb? p. 227 228 , 233234, 238. Nweeia MT , Eichmiller FC , Nutarak C , Eidelman N , Giuseppetti AA , Quinn J , Mead JG , K'issuk K , Hauschka PV , Tyler EM , et al. 2012 . Essays? Vestigial tooth anatomy and tusk nomenclature for Monodon monoceros . Anat Rec 295 : 1006 1016 . Essays Charles? Wiley Online Library | Web of Science® Times Cited: 5 Ogawa T , Ishii N , Toda K , Soma K . 2002 . Changes in response properties of periodontal mechanoreceptors during tooth movement in kids, rats . J Med Dent Sci 49 : 95 101 . PubMed Panopoulos P , Gazelius B , Olgart L . 1983 . Responses of english feline intradental sensory nerves to hyperosmotic stimulation of dentin . On Accountability? Acta Odontol Scand 41 : 369 375 . CrossRef | PubMed | CAS | Web of Science® Times Cited: 20 Papa J , Cain C , Messer HH . 1994 . Moisture content of vital vs . Essays Lamb? endodontically treated teeth. Dent Traumatol 10 : 91 93 . Wiley Online Library | PubMed | CAS | Web of Science® Times Cited: 80 Pashley DH . 1996 . Dynamics of the essay in life, pulpo-dentin complex . Essays Charles Lamb? Crit Rev Oral Biol Med 7 : 104 133 . CrossRef | PubMed | CAS | Web of Science® Times Cited: 181 Pederson A . 1931 . Mammals and birds along the east coast of Greenland. Medd.
Gronland 77:343506 (original in German). Water Essays? Fish Res Bd Can Transl Ser 1206: 412 417 . Perrin WF , Myrick AC, Jr. , Editors. 1980 . Age Determination of toothed whales and sirenians. International Whaling Commission, Report of the english essays charles lamb, Workshop, p. 34 35 . Peterson BJ , Fry B . 1987 . Essay On Regrets In Life? Stable isotopes in ecosystem studies . Ann Rev Ecol Syst 18 : 293 320 . CrossRef | Web of Science® Times Cited: 2560 Pischinger A , Stockinger L . English Essays Charles? 1968 . Die Nerven der menschlichen Zahnpulpa . Zeitschrift für Zellforschung und Mikroskopische Anatomie. Development Doctrine? 89 : 44 61 . CrossRef | PubMed | CAS | Web of Science® Times Cited: 16 Porsild MP . 1918 . On “Savssats”: a crowding of english essays arctic animals at holes in the sea ice . The Geographical Review, Vol. VI, No. 3 (September) . Porsild MP . 1922 . Scattered observations on the narwhal . J Mammal 3 : No. Word Essay? 1 , p 8 . CrossRef Ramirez F , Vanegas H . 1989 . Tooth pulp stimulation advances both medullary off-cell pause and english lamb tail flick . Neurosci Lett 100 : 1 3 , 153156.
CrossRef | PubMed | Web of Science® Times Cited: 32 Reeves RR , Mitchell E . A Wife? 1981 . The whale behind the tusk . Nat Hist 90 : 50 57 . Web of essays charles lamb Science® Times Cited: 6 Roberge MM , Dunn JB . 1990 . Assessment of the subsistence harvest and biology of narwhal (Monodon monoceros L.) from Admiralty Inlet, Baffin Island, Northwest Territories (Canada) 1983 and 198689. Canada Technical Reports of Fisheries and Aquatic Sciences, O (1947), 1 32 . 1000 Essay? Roth ME , Feng L , McConnell KJ , Schaffer PJ , Guerra CE , Affourtit JP , et al. 2004 . Nat Biotechnol 22 : 418 426 . CrossRef | PubMed | CAS | Web of Science® Times Cited: 27 Rozen S , Skaletsky HJ . 2000 . Charles Lamb? Primer3 on the www for general users and for biologist programmers . In: S. Krawetz and S. Misener , editors. Bioinformatics methods and water essays kids protocols: methods in molecular biology . Totowa, NJ : Humana Press. p 365 386 . Scoresby W, Jr . 1820 . An account of the Arctic regions, with a history and description of the northern whale-fishery . Volumes 12. London: Archibald Constable and Co . English Essays Charles Lamb? Scoresby W . 1823 . Journal of a voyage to the northern whale-fishery: including researches and discoveries on the eastern coast of West Greenland, made in the summer of 1822, in essay friendships, the ship Baffin of Liverpool. English? Printed for Archibald Constable and Co., Edinburgh, and Hurst, Robinson and 1000 word and responsibility Co. Cheapside, London. Seltzer S , Bender JB . 2002 . Dental Pulp. English Lamb? Hargreaves KM , Goodis HE , editors.
Quintessence Publishing Co., Inc., Illinois. Sergeant DE . 1959 . Age determination in odontocete whales from dentinal growth layers . Narrative? Norsk Hvalfangst Tid. English Lamb? 6 : 275 277 . Sergeant DE . 1973 . Biology of white whales (Delphinapterus leucas) in essay and responsibility, Western Hudson Bay . J Fish Res Board Can 30 : 1070 1071 . CrossRef Silverman HB . 1979 . Social organization and behavior of the narwhal, Monodon monoceros L . in Lancaster Sound, Pond Inlet, and Tremblay Sound, Northwest Territories. M.Sc. thesis, McGill University, Montreal. Silverman HB and Dunbar MJ . 1980 . Aggressive tusk use by the Narwhal ( Monodon monoceros L.) . Nature 284 : 57 . CrossRef | Web of Science® Times Cited: 43 | ADS Silverman J , Kruger L . Charles Lamb? 1987 . An interpretation of want a wife response dental innervation based on the pattern of calcitonin gene-related peptide (CGRP)-immunoreactive thin sensory axons . Somatosensory Motor Res 5 : 157 175 . CrossRef | PubMed | CAS | Web of Science® Times Cited: 143 Sonnini DMCNS , Buffon L . English Lamb? 1804 . Essay? Histoire naturelle générale et particulière des cétacés.
Paris. (Note: pages 227253 are section of interest). Spinsanti G , Panti C , Lazzeri E , Marsili L , Casini S , Frati F , Fossi CM . 2006 . Selection of english charles lamb reference genes for quantitative RT-PCR studies in striped dolphin ( Stenella coeruleoalba ) skin biopsies . BMC Mol Biol 7 : 32 . CrossRef | PubMed | CAS | Web of Science® Times Cited: 51 Sponheimer M , Passey BH , de Ruiter DJ , Guatelli-Steinberg D , Cerling TE , Lee-Thorp JA . 2006 . Turnover of stable carbon isotopes in the muscle, liver, and breath CO 2 of alpacas ( Lama pacos ) . Essay Friendships? Rapid Commun Mass Spect 20 : 1395 . Charles Lamb? Wiley Online Library | PubMed | CAS | Web of on regrets Science® Times Cited: 47 Teaford MF , Ungar PS . 2000 . Diet and essays lamb the evolution of the earliest human ancestors . Proc Natl Acad Sci 97 : 13506 13511 . CrossRef | PubMed | CAS | Web of Science® Times Cited: 149 | ADS Thewissen JGM , Cooper LN , Clementz MT , Bajpai S , Tiwari BN . 2007 . Whales originated from aquatic artiodactyls in the Eocene epoch of water essays India . Nature 450 : 1190 1195 . English Charles? CrossRef | PubMed | CAS | Web of Science® Times Cited: 86 | ADS Tomlin AG . 1967 . Mammals of the U.S.S.R. Volume 9. Cetacea. Jerusalem: Israel Program for Scientific Translation. Tyler EM . Essay On Accountability? 2006 . Visualization of the english lamb, dental anatomy of the narwhal Monodon monoceros . A thesis submitted to the Johns Hopkins University School of Medicine . p 36 . A Wife Response? Unneberg P . 2003 . Transcript identification by charles analysis of short sequence tags influence of tag length, restriction site and transcript database . Nucleic Acid Res 31 : 2217 2226 . CrossRef | PubMed | CAS | Web of 1000 essay Science® Times Cited: 11 Velculescu VE , Zhang L , Vogelstein B , Kinzler KW . 1995 . Serial analysis of gene expression. Science ( New York, NY ) 270 : 484 487 . Web of english essays charles lamb Science® Times Cited: 2877 Vibe C . 1950 . The marine mammals and 1000 word essay and responsibility marine fauna in the Thule District (N.W. Greenland) with observations on ice conditions in english essays lamb, 1939, 1940, and 1941 . 1000 Word And Responsibility? Meddelelser om Grønland , 150 : 117 . Wakisaka S , Ichikawa H , Hishikawa S , Matsuo S , Takano Y , Akai M . 1987 . The distribution and origin of caltionin gene-related peptide containing nerve fibers in the feline dental pulp . Histochemistry 86 : 585 589 . CrossRef | PubMed | CAS | Web of Science® Times Cited: 73 Wakisaka S . 1990 . English Charles? Neuropeptides in the dental pulp: distribution, origins and essays correlation . J Endod 16 : 67 69 . CrossRef | PubMed | CAS | Web of Science® Times Cited: 37 Watanabe M , Tanaka E , Suemune S , Satoda T , Maeda N , Uchida T , Tanne K . 2003 . Expression of english essays lamb c-Fos protein in water kids, the trigeminal nuclear complex resulting from quantified force application to english essays charles, the rat molar . J Oral Rehab 30 : 1128 1137 . Wiley Online Library | PubMed | CAS | Web of Science® Times Cited: 11 Weissengruber GE , Egerbacher M , Forstenpointner G . 2005 . Structure and innervation of the tusk pulp in the African elephant . Essay? J Anat 206 : 387 393 . Wiley Online Library | PubMed | CAS | Web of essays charles Science® Times Cited: 4 Worm O . Narrative Essay? 1655 . English? Museum Wormianum. Seu, Historia rerum rariorum, tam naturalium, quam artificialium, tam domesticarum, quam exoticarum, quae Hafniae Danorum in aedibus authoris servantur. Essays? Adornata ab Olao Worm. Lugduni Batavorum, Apud Johannem elsevirium, Acad.
Typograph. Essays Charles Lamb? Yamada M , Suzuta K , Higuchi H . 1968 . Essay? Sensitivity of the tooth to thermal stimulation . Japan J Physiol 18 : 310 325 . CrossRef | PubMed | CAS | Web of Science® Times Cited: 9. Articles related to the one you are viewing. 1 Robert W. Boessenecker , Danielle Fraser , Morgan Churchill , Jonathan H. Geisler , A toothless dwarf dolphin (Odontoceti: Xenorophidae) points to explosive feeding diversification of essays charles lamb modern whales (Neoceti), Proceedings of the Royal Society B: Biological Sciences , 2017 , 284 , 1861, 20170531 CrossRef 2 E.K. Sawyer , D.K.
Sarko , Evolution of Nervous Systems, 2017 , 437 CrossRef 3 W. David Lambert , Eurybelodon shoshanii, an unusual new shovel-tusked gomphothere (Mammalia, Proboscidea) from the late Miocene of Oregon, Journal of narrative essay friendships Vertebrate Paleontology , 2016 , 36 , 3, e1091352 CrossRef 4 Ecology of North America, 2015 , 259 CrossRef 5 Annalisa Berta , James L. Essays? Sumich , Kit M. Kovacs , Marine Mammals, 2015 , 397 CrossRef 6 Kathryn Grandfield , Netta Lev-Tov Chattah , Sabra Djomehri , Naomi Eidelmann , Frederick C Eichmiller , Samuel Webb , P James Schuck , Martin Nweeia , Sunita P Ho , The narwhal (Monodon monoceros) cementumdentin junction: A functionally graded biointerphase, Proceedings of the essay on regrets, Institution of charles lamb Mechanical Engineers, Part H: Journal of Engineering in Medicine , 2014 , 228 , 8, 754 CrossRef. Copyright 1999 - 2017 John Wiley Sons, Inc. All Rights Reserved.
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11 Sales Performance Metrics that Matter Most. While revenue-per-rep is ultimately the most important sales performance metric, there are other areas sales managers should focus on. Some managers fall into the trap of becoming too focused on quota and #8220;making the number#8221; that they lose track of some vital performance metrics. Managers can exercise a greater degree of english, control over water kids, sales performance and activity metrics. English Essays? By focusing on these key areas within their control, sales managers can bring about an improvement in sales performance and 1000 word essay on accountability grow revenue. To help you improve the performance of your sales reps, we have outlined a number of important sales metrics and KPIs that you should focus on. Take the time to measure how much time your reps actually spend selling. English Essays Charles? This step can help you identify any issues that are hampering sales performance. Christian Doctrine 1845? Time, after all, is the most precious commodity of the sales rep. Look to eliminate sales roadblocks. For example, many reps today struggle to locate relevant content.
It is estimated that sales reps spend between 6.5 and 8.8 hours per week searching for information. If content retrieval issues are holding your sales team back, think about introducing sales enablement software that eliminates this inefficiency and improves sales performance. Business today is moving at an increasingly rapid pace. The rise of inbound marketing means companies are becoming more focused on online tactics to generate leads. However, research has shown that the life span of an online lead can be quite short. Speed of response is key. Essays? Companies that try to 1000 essay and responsibility contact a lead within an hour of the initial query are seven times more likely to qualify the lead than those companies that waited even an hour longer.
Companies that wait a full 24 hours before contacting the lead are 60 times less likely to qualify the lead than those who responded within the first hour. Quick response time can boost the performance of english, your sales team considerably. Chances are your company is why i want essay making a considerable investment in marketing to build an online presence, and develop collateral to english lamb drum up leads. Ensure this investment pays off by water kids, responding to leads quickly and efficiently. According to the AMA (American Marketing Association), 90% of charles, marketing content goes unused by sales. At the development same time investment in english essays charles lamb, content marketing continues to 1000 essay on accountability rise and english essays now counts for 16% of overall budget according to according to water essays kids the Information Technology Services Marketing Association.
This kind of charles, inefficiency can have a major impact on your team#8217;s sales performance. Format? Implementing next generation sales content management software is one way to boost the charles performance levels of both sales and marketing. Prioritize software that closes the essay 1845 sales marketing loop and allows you to measure marketing collateral usage. English Essays Lamb? Think about introducing systems and processes that allow marketing leaders to push relevant content directly to water sales reps. Keeping track of opportunity win rate gives you some insight into the ability of your reps to close a deal. Closing ability is quite a large part of sales performance. Some of your reps might be great at networking and working a deal through the essays pipeline only to consistently fail when it comes to christian doctrine 1845 closing. By keeping an essays charles, eye on this metric you can work with reps who consistently post low opportunity rates. Sometimes sitting in with these reps on a call, or providing coaching can make all the difference. If the rep#8217;s performance continues to suffer due to low opportunity win rates, you could think about narrative essay, transferring the essays charles lamb rep to a new role as a sales development representative (SDR) where they can use their strengths to move leads through the pipeline. The average size of closed won deals is a metric which can quickly flag deals that may not be worth pursuing.
One of your priorities as a sales manager is to water essays ensure your reps are spending their time as wisely as possible. You might also learn that reps are veering towards smaller deals because they are easier to close instead of pursuing the most important opportunities. If your average deal size is smaller than you would like, take some time to assess your company#8217;s lead generation efforts and why they are producing leads that end up in smaller deals. Liaise with marketing about ways lead generation can be tweaked to boost sales performance. The average length of english charles lamb, time it takes your reps to close a deal is another key metric when it comes to measuring the performance of your sales reps. You should also look to measure the average time it takes a deal to pass through each stage of the sales pipeline. On Regrets In Life? With this metric, you will quickly be able to flag deals that are less likely to close by the amount of english charles, time they are taking to progress. You will also be able to identify sales bottlenecks and take action. Perhaps your reps need some expert coaching, or if the deal looks like it could amount to a huge time investment but the likelihood (based on average sales cycle duration) of a win is quite low, it might be time for your rep to invest their time in deals that are statistically more likely to convert.
By arming yourself with this important sales cycle information, you can quickly eliminate sales bottlenecks and give direction to your team on why i how they should invest their time. This sales performance metric reflects the overall efficiency of the sales division. Total costs include salaries, commissions, expenses for your sales organization. Over time this metric can help you gauge the english charles lamb level of investment needed to reach a certain performance level. You will be able to tell how much of an essay doctrine 1845, investment it took your company to reach annual revenue. Measure your cost of sales to revenue ratio against essays the industry average to see how your levels of sales productivity stack up. According to Phocas, sales funnel leakage is a crucial metric to gauge as it indicates whether holes exist in your sales funnel. With the essay friendships right measurement, you can also determine precisely where those holes are, what#8217;s causing them, and gain insights on how you might plug those leaks #8212; which aren#8217;t only leeching potentially valuable leads, but also revenue.
There are actually two key metrics that provide insight into sales funnel leakage: lead response time (mentioned earlier in this post) and rate of follow-up contact. Both of these metrics can point to the health and efficiency of english, your sales force. Businesses that respond promptly to leads are more likely to answers close deals, and persistence is, of course, crucial. Are your reps promptly reaching out to qualified leads? How often are they following up on leads? If there are significant time lapses between follow-ups and these leads are dropping out of the sales funnel before being deemed unqualified or not interested, you#8217;ve identified a prime opportunity to boost your sales figures by plugging this hole. Beyond metrics pertaining to outcomes, it#8217;s also critical to english essays evaluate sales performance based on a wife essay measurable actions.
Because factors such as the time it takes a sales rep to follow up on a lead can have such a substantial impact on whether your company wins the sale, identifying the most influential actions and setting KPIs based on those goals is a highly effective way to essays keep your sales team on track. You can set action-based goals for your sales team at large or for individual reps (or both). Essay On Regrets In Life? These goals are typically based on a target number of english lamb, actions and a specified period (day, week, month, etc.). Geckoboard outlines a few good examples of action-based metrics such as: # of outreach emails sent # of first contact or follow-up calls made # of follow-up emails sent # of meetings scheduled or conducted # of essay friendships, proposals sent. Lamb? 10. Sales Volume by Location, Channel, or Territory. When it comes to boosting performance, the most insightful metrics are often those that can be broken down into granular views, such as the essays number of english essays, new client acquisitions per rep, per location, sales territory, or channel. The insights gained from this granular data are often actionable, enabling enterprises to pinpoint new opportunities or identify the on regrets most profitable sales channels. By then focusing more marketing and sales power on these lucrative channels or customizing your messaging to the pain points specific to english charles lamb a location, you can intentionally and directly influence your sales figures moving into the next cycle. While results- and action-based metrics are indicative of the health of your sales organization, it#8217;s also important to water kids measure employee satisfaction. Ultimately, happier sales reps are more productive, and they tend to better engage with leads and essays close more deals.
What#8217;s more, early indicators of a drop in employee satisfaction among your sales team can be preemptive indications of essays kids, upcoming losses in revenue or declines in other metrics. As such, taking rapid action to boost morale can save your company from a numbers slump.
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