Giraffe

Giraffes emerged during the Oligocene (34–23million years ago) when they last shared a common ancestor with their closest living relative, the okapi, which is the only other living member of the tribe Giraffini (Hassanin and Douzery, 2003).

From: Evolution of Nervous Systems (Second Edition) , 2017

Conservation Status of Giraffe: Evaluating Contemporary Distribution and Abundance with Evolving Taxonomic Perspectives

Michael B. Brown , ... Julian Fennessy , in Reference Module in Earth Systems and Environmental Sciences, 2021

Abstract

Giraffe are iconic figures across a range of African landscapes but they are currently under considerable conservation threat. Although they are widely distributed throughout 21 different countries, continent-wide populations have declined considerably over the past several decades, highlighted by the International Union for the Conservation of Nature's (IUCN) new categorization of giraffe as a single species as " Vulnerable." Recent genetic studies, however, propose alternative taxonomic categorizations in which giraffe are comprised of four distinct species. These proposed taxonomic classifications have considerable impact on giraffe conservation status, emphasizing the diverse challenges that giraffe face throughout Africa. Here, we describe recent studies on the taxonomic status of giraffe and examine implications for conservations status assessments. We conducted an extensive review of current giraffe abundance throughout all known populations and evaluated these updated abundance trends through the taxonomic perspective of a four species classification. We provide the most recent and comprehensive abundance estimates for wild giraffe in Africa. According to our assessment, there are approximately 117,173 giraffe in the wild. Providing the most current and accurate giraffe abundance estimates within evolving taxonomic perspectives can better guide targeted conservation efforts for these imperiled taxa.

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Giraffe Demography and Population Ecology

D.E. Lee , M.K.L Strauss , in Reference Module in Earth Systems and Environmental Sciences, 2016

Giraffe Life History

Giraffes are endemic African ruminants ( Fig. 1 ) that are long-lived (Jolly, 2000; King, 1947), have delayed maturity (Hall-Martin and Skinner, 1978), and cannot reproduce every year due to a gestation length of 14.7 months (del Castillo et al., 2005), so we suggest the following modifications to the impala population model described earlier. We will maintain the 1-year time increment.

Fig. 1. Current range of giraffe (G. camelopardalis) subspecies.

 http://maps.iucnredlist.org/map.html?id=9194 © International Union for Conservation of Nature and Natural Resources.

Annual probability of survival varies considerably among age classes (Foster and Dagg, 1972), so survival probabilities will differ for juveniles (calves or young of the year), subadults, and adults (Lee et al., 2016). Individuals show variation in the age at which they become an adult and first breed, and we formulate this parameter as the probability of first breeding, which we symbolize as β (Nur and Sydeman, 1999). β is age-specific like survival and RS (Lee et al., 2012). At a young age, β  =   0, minimum age of first breeding corresponds to the youngest age at which β  >   0.

"Reproductive success" can be separated into two components: (1) the probability that an experienced adult attempts to produce offspring and (2) RS among those breeding individuals. The first component is "breeding propensity" symbolized by γ. The second component is then the number of live calves produced per female that attempted breeding that year, which accounts for lost pregnancies and stillborn calves. RS can also be formulated as a combined probability, in which case it is the number of calves per adult female per year (for more detail, see section " Reproductive Success ").

Movement probabilities are also likely to be age-specific, especially for dispersing subadults, but this is a topic with very little data at present. We assume calves move with their mothers during their first year of life. For more detail, see section " Movement: Immigration and Emigration ."

Thus, we can parameterize giraffe demographic processes determining population growth in terms of eight parameters: (1) adult survival, (2) subadult survival, (3) juvenile survival, (4) recruitment probability, (5) breeding propensity, (6) RS per breeder, (7) adult net migration (with their calves), and (8) subadult net migration.

This formulation developed for giraffes is also appropriate for other long-lived herbivores. One peculiarity of giraffes and many other tropical species is that conceptions and births can occur in every month of the year (Zerbe et al., 2012), so a birth-flow reproduction model is required (Caswell, 2001; Caughley, 1977). Many different parameterizations of population dynamics are possible, but we believe this is both comprehensive and tractable.

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In Loeffler's Footsteps – Viral Genomics in the Era of High-Throughput Sequencing

Sandra Blome , ... Kerstin Wernike , in Advances in Virus Research, 2017

3.1 Giraffe Virus

The Giraffe pestivirus was associated with an outbreak of MD-like symptoms in giraffes in the Nanyuki District of Kenya in 1967 ( Dekker et al., 1995; Harasawa et al., 2000). The virus was shown to be divergent from classical pestiviruses (Becher et al., 1997, 2003). A closely related virus, referred to as PG-2, was obtained from a bovine cell culture in the 1990th (Becher et al., 2003) suggesting the presence of further, similar viruses in Africa. The knowledge about the natural host range, virulence, or geographic distribution is scarce until now; however, the transmission of such newly emerging pestiviruses to domestic ruminants could pose a risk for the livestock industry, especially in countries like Switzerland, Denmark, Sweden, or Germany with eradication programs and more and more naïve cattle (Moennig and Becher, 2015; Moennig et al., 2005; Stahl and Alenius, 2012).

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Horns and Ossicones

Brian K. Hall , in Bones and Cartilage, 2005

Giraffes

Giraffes ( Giraffa camelopardalis), as the tallest animals in the world, are among the most well known of all mammals. A male standing close to 5.5 metres (16–18 feet), weighing almost 2000 kg and consuming over 60 kg of leaves per day is an amazing sight in the wild and a challenge to keep in captivity. Nine subspecies are recognized, the reticulated giraffe, Giraffa camelopardalis reticulata, and Rothschild's giraffe, Giraffa camelopardalis rothschildi, being the most frequent subspecies seen in zoos. The specific epithet comes from cameleopard, one of the early names for this animal thought to be part camel and part leopard. The Chinese named it 'K'i-lin' after a mythical animal resembling a unicorn. In the Koran it is the 'serafe' or lovely one.

The paired structures on the heads of giraffes and okapi are neither horns nor antlers, but ossicones, sometimes referred to as knobs. Ossicones, permanent bony outgrowths covered with vascularized skin, have fascinated biologists ever since Richard Owen (1841) published his paper on the anatomy of the Nubian giraffe, Giraffa camelopardali typica. Lydekker (1904) thought he could identify subspecies in part based on the horns.

According to Ganey et al. (1990) ossicones are not true horns because they lack a keratin sheath. Another complication when making comparisons is that giraffes have several sets of ossicones, the most prominent of which are those on the parietal bones, but with, in addition, a more anterior median horn on the frontal bone of all males and a few females – antlers develop on the frontal bones – an occipital ossicone on the occipital bone, orbital ossicones associated with the eyes, and an azygous ossicone just posterior to one of the orbital ossicones. Sexual dimorphism in ossicones relates to the mode of fighting; males use their heads as clubs, females kick with their legs. Constant awareness of their environment is achieved by sleeping for no more than half an an hour a day, the half hour spread over some five-minute 'naps' throughout the day.

Ossicones begin as nodules of bone beneath the skin, quite separate from the bone of the skull – like the human patella, they are sesamoid bones – but each ossicone later fuses with the underlying bone. The overlying skin is not shed, in contrast to the annual shedding of the velvet from antlers. Again, in contrast to antlers, ossicones are permanent and present in both sexes; in the Miocene moose-like Asian and African giraffids, Sivatherium giganteum and S. maurusium, the ossicones (horns) were almost a metre in length and palmate.

In one of the few recent studies, Spinage (1968) claimed that ossicones are present in foetal giraffes as cartilaginous precursors, basing this claim on Ray Lankester's study published in 1907. The line drawing Spinage provides as Figure 4 is not informative, although he describes ossification of the cartilage commencing after birth, beginning apically and progressing proximally, leaving a pad of cartilage at the base for subsequent growth – the latter based on Blumenbach's 1805 Handbuch and an 1872 paper by Murie, both of whom thought the ossicones developed endochondrally from this cartilage.

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The Nervous Systems of Early Mammals and Their Evolution

M.A. Raghanti , ... P.R. Hof , in Evolution of Nervous Systems (Second Edition), 2017

2.14.3.3 Giraffes

As mentioned previously, our knowledge of giraffe neuroanatomy is limited to a handful of studies. A systematic analysis of neuron morphologies revealed the expected agranular organization for motor cortex, but a granular visual cortex, unlike hippopotamids and cetaceans ( Butti et al., 2014, 2011; Hof et al., 2005; Hof and Van der Gucht, 2007; Jacobs et al., 2015, 1984), but giraffes do not possess the expanded layer IV in the visual cortex that characterizes primate species.

Overall, giraffe neuron morphology was similar to what has been described for other cetartiodactyls (Butti et al., 2015, 2011; Jacobs et al., 2015). However, the giraffe was unique from other species in having a variety of complex neocortical spiny neurons that possess either single or narrowly bifurcating apical dendrites that are arrayed in a columnar fashion and contribute to the minicolumns of the neocortex (Jacobs et al., 2015). Spiny neurons included the typical pyramidal morphologies (ie, magnopyramidal and extraverted neurons in motor and visual cortex and gigantopyramidal neurons in motor cortex), pyramidal neurons that were horizontally oriented, and unusual crablike neurons that are bitufted with horizontally projecting dendrites located in the visual cortex (Jacobs et al., 2015, 2014, 2016). Extraverted neurons, located closer to the pial surface, were the second most common neuron type and are common across species that possess agranular cortex. Giraffe pyramidal neurons bear a close resemblance to those found in pig, sheep, and cetaceans but differ from those of horse, cow, and elephants in that the latter group possesses widely bifurcating, V-shaped apical dendrites, and the former possess prominent apical dendrites that bifurcate some distance from the cell body (Jacobs et al., 2015). However, the cetacean-specific tritufted pyramidal neuron and Sternzellen were not observed in giraffe cortex. Giraffe magnopyramidal neurons were similar to the magnopyramidal neurons of pygmy hippopotamus motor cortex and the gigantopyramidal neurons observed in pig. The giraffe magnopyramidal neurons that were observed in layers V and VI are common among laurasiatherians (ie, mammals that are thought to have originated in Laurasia and include odd- and even-toed ungulates, bats, shrews, carnivores, and whales) and their occurrence in visual cortex indicates that vision is important in this species (Coimbra et al., 2013; Jacobs et al., 2015). As for the atypical spiny neuron types in giraffe cortex, the horizontally oriented spiny neurons are also present in a variety of other species, including carnivores, cetartiodactyls, and primates while the crablike neurons are absent in cetaceans. Aspiny cortical neurons were more uniform with a morphology that is conserved across eutherian mammals (Jacobs et al., 2015). Interestingly, neuron morphologies in the newborn giraffe were qualitatively similar to those of adults (Jacobs et al., 2016). Newborn interneurons were of a similar size to adult interneurons whereas projection neurons were smaller in the newborn. Local circuit neurons were spiny, in contrast to the adult aspiny forms. Both newborn and adult giraffe neuron morphologies are consistent with what has been reported for other cetartiodactyls.

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Basic Sleep Concepts, Science, Deprivation, and Mechanisms

J. Siegel , in Encyclopedia of Sleep, 2013

Sleep Amounts in Mammals and Birds

The sleep durations of animals vary widely. Some animals such as the giraffe sleep as little as 2  h a day. Others such as the opossum sleep 18   h a day. Humans typically have 7–8   h of sleep a day with 4   h in REM sleep. It was hypothesized that differences in sleep across species were correlated with some physiological feature of each animal, such as body mass, brain size, intelligence, body temperature, or life span. If so, this would have been a starting point for determining the physiological function of sleep. However, although some weak relations were found across mammals, these relationships mostly evaporated as the collected data and analysis techniques increased in quality and amount. Significant relations explained only a very small portion of the variance and even these relations appear to be largely a function of the mathematical treatment of the data. Measures that attempt to factor in sleep 'intensity' to explain the variability of sleep duration across species do not appear to solve this problem, as animals that sleep longer also appear to sleep more deeply.

A further problem with this approach is that to the extent that very weak and variable relations between the above physiological variable and sleep state have been claimed in mammals, none of these relations appears to hold in birds. Thus, this work does not appear to be leading to convincing evidence for a simple physiological explanation for sleep time or sleep stage duration across species ( Figure 1 ).

Figure 1. Sleep time is not strongly related to physiological variables. Examples of reported mammalian and bird sleep amounts are shown (see Siegel 2001, 2005, 2008, 2009 for further examples). Noteworthy is the similarity of sleep parameters in the guinea pig and baboon, animals that dramatically differ in almost every physiological parameter, and the very high level of sleep in the big brown bat and the variability of REM sleep parameters in birds. A variety of studies have attempted to correlate total sleep times and REM sleep times or percentages with physiological variables, for example, metabolism, life span, brain/body weight ratio, and altricial–precocial status. (Early studies concluded that animals that were born in a relatively immature, helpless state (altricial) had more REM sleep as adults; however, later studies have disputed this.) Recent studies have found that relations between physiological variables and sleep duration disappear or reverse depending on which animals are included and excluded and how the data are handled. The few studies finding significant relations between sleep and physiological variables have found correlations that explain very little of the enormous variance in sleep parameters across homeotherms (Siegel, 2009).

 Credits for pig and baboon in Siegel 2001; Bat, billbatboy ca.; Owl, Kim Taylor; Zebra finch, central pet; Mallard, U Michigan Zoology.

An important recent development has been the appreciation of the differences between land mammals and marine mammals. Dolphins never show the long periods of bilaterally symmetrical high-voltage EEG activity that are used for identification of non-REM sleep in land mammals. Rather one hemisphere exhibits a sleep-like EEG while the other exhibits a wake-like EEG. Convincing evidence for REM sleep has not been seen in these marine mammals. Other subsequently studied cetacean species have shown the same unihemispheric sleep waves and apparent lack of REM sleep. The animals may be mobile and responsive during this EEG pattern, making tenuous any assumption that this is 'sleep,' of the kind that is seen in most land mammals studied. This asymmetrical EEG pattern may be related to the apparent lack of REM sleep in these mammals.

Studies in land mammals have generally shown that REM sleep amounts are greatest at birth. Therefore it was thought that REM sleep in dolphins might be observed if they were examined shortly after birth. However, when this was done, it was surprising to find that, unlike land mammals, both newborn dolphins and their mothers were continuously active 24   h day for weeks after birth. Similar observations were made in killer whales. In the wild, the postpartum period is characterized by migration over long distances from birthing grounds to feeding grounds. Dolphins and even killer whales are subject to predation at these times, so sensorimotor responsiveness is even more important for these species during migration than when they are in familiar waters. Although further studies are necessary, all indications are that they migrate while in a highly aroused state, rather than a 'sleep swimming' state. The healthy fur seal and walrus, under controlled ad libitum feeding conditions, will often spontaneously stay awake for 24–48-h periods, as indicated by both behavioral and EEG observations, a behavior only rarely observed in land mammals.

Interesting observations in the white crowned sparrow, a bird that migrates long distances, produced similar results. When these birds are kept caged in the laboratory they show greatly reduced sleep and increased activity during the periods when they would be migrating in the wild. This period of greatly reduced sleep and greatly increased activity and wing flapping lasts for several weeks. Despite this, there is no increased sleep after this period to 'make up' for lost sleep. This was also seen in observations of cetaceans. It can be shown that lack of rebound sleep was also seen after the period of postpartum activity in cetaceans.

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Evolution of sleep (sleep phylogeny)

Jerome Siegel , in Reference Module in Neuroscience and Biobehavioral Psychology, 2021

Sleep amounts in mammals and birds

The sleep durations of animals vary widely. Some animals such as the giraffe and elephant sleep as little as 2  h a day. Others such as the opossum sleep 18   h a day (Gravett et al., 2017; Siegel, 2001, 2005, 2008, 2009b, 2021b; Zepelin et al., 2005). Humans typically have 7–8   h of sleep/day with 2   h in REM sleep. It was hypothesized that differences in sleep across species were correlated with some physiological feature of each animal, such as body mass, brain size or intelligence, body temperature, lifespan etc. If so, we would have a starting point for determining the physiological function of sleep. However, although some weak relations were found across mammals, as the collected data and analysis techniques increased in quality and amount, these relationships have largely evaporated (Siegel, 2008, 2009b). Significant relations explained only a very small portion of the variance and even these relations appear to be largely a function of the mathematical treatment of the data (Fig. 1). Measures that attempt to factor in sleep "intensity" to explain the variability of sleep duration across species do not appear to solve this problem, since animals that sleep longer appear to sleep more deeply as well (Siegel, 2001, 2005, 2008, 2009b, 2021a).

Fig. 1. Sleep time is not strongly related to physiological variables. Examples of reported mammalian and bird sleep amounts are shown (see Siegel 2001, 2005, 2008 and 2009a,b for further examples). Noteworthy is the similarity of sleep parameters in the guinea pig and baboon, animals that dramatically differ in almost every physiological parameter, and the very high level of sleep in the big brown bat and the variability of REM sleep parameters in birds. A variety of studies have attempted to correlate total sleep times and REM sleep times or percentages with physiological variables, e.g., metabolism, lifespan, brain/body weight ratio, altricial-precocial status. (Early studies concluded that animals that were born in a relatively immature, helpless state (altricial) had more REM sleep as adults, however later studies have disputed this). Recent studies have found that relations between physiological variables and sleep duration disappear or reverse depending on which animals are included and excluded and how the data are handled. The few studies finding significant relations between sleep and physiological variables have found correlations that explain very little of the enormous variance in sleep parameters across homeotherms (Siegel, 2009a,b).

 Credits for pig and baboon in Siegel (2001); Bat, billbatboy.ca; Owl, Kim Taylor; Zebra finch, central pet; Mallard, U Michigan Zoology.

A further problem with this approach is that to the extent that very weak and variable relations between the above physiological variables and sleep states have been claimed in mammals, none of these relations appears to hold in birds (Roth et al., 2006). Thus, this work does not appear to be leading to convincing evidence for a simple physiological explanation for sleep time or sleep stage duration across species.

An important recent development has been the appreciation of the differences between land mammals and marine mammals. Dolphins never show the long periods of bilaterally symmetrical high voltage EEG activity that are used for identification of nonREM sleep in land mammals. Rather one hemisphere exhibits a sleep like EEG while the other exhibits a wake like EEG. Convincing evidence for REM sleep has not been seen in these marine mammals. Other subsequently studied cetaceans species have shown the same unihemispheric sleep waves and apparent lack of REM sleep (Mukhametov, 1984). The animals may be mobile and responsive during this EEG pattern, making tenuous any assumption that this is "sleep," like that seen in most studied land mammals (Siegel, 2008). This asymmetrical EEG pattern may be related to the apparent lack of REM sleep in these mammals (Lyamin et al., 2008c; Siegel, 2008, 2009b). Fur seals have been found to have sleep amounts resembling those of humans when on land. But they sleep "unihemispherically" like dolphins when they are at sea, when they spend most of their lives (Lapierre et al., 2007, 2013a,b). Most strikingly, the evidence suggests that during the 7 months of the year in which they are at sea, they have little or no REM sleep (Lyamin et al., 2018; Siegel, 2021a).

Studies in land mammals have generally shown that REM sleep amounts are greatest at birth. Therefore we thought we might observe REM sleep in dolphins if we examined them shortly after birth. However, when we did this we were surprised to find that unlike land mammals, both newborn dolphins and their mothers were continuously active 24   h/day for weeks after birth. We made similar observations in killer whales (Lyamin et al., 2005). In the wild, the postpartum period is characterized by migration over long distances from birthing grounds to feeding grounds. Dolphins and even killer whales are subject to predation at these times, so sensory-motor responsiveness is even more important for these species during migration than when they are in familiar waters. Although further studies are necessary, all indications are that they migrate while in a highly aroused state, rather than a "sleep swimming" state. The healthy fur seal and walrus, under controlled ad libitum feeding conditions, will often spontaneously stay awake for 24–48   h periods, as indicated by both behavioral and EEG observations, a behavior only rarely observed in land mammals (Lyamin et al., 2008a,b; Pryaslova et al., 2009).

Beautiful observations in the white crowned sparrow, a bird that migrates long distances produced similar results (Rattenborg et al., 2004). When these birds are kept caged in the laboratory they show greatly reduced sleep and increased activity during the periods when they would be migrating in the wild. This period of greatly reduced sleep and greatly increased activity and wing flapping lasts for several weeks. Despite this, there is no increased sleep after this period to "make up" for lost sleep. This was also seen in our observations of cetaceans. It can be shown that the learning and performance abilities of the white crowned sparrow increase during this sleepless migratory period, which has been compared to manic periods in humans (Rattenborg et al., 2004).

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The Nervous Systems of Early Mammals and Their Evolution

E.K. Sawyer , D.K. Sarko , in Evolution of Nervous Systems (Second Edition), 2017

2.22.2.6 Major Clade Laurasiatheria, Order Artiodactyla

The artiodactyls are the even-toed ungulates, which include the hippopotamuses, deer, giraffes, llamas, camels, pigs, cows, sheep, and goats, among others. Modern phylogenies also put cetaceans in this order ( Graur and Higgins 1994; Murphy et al., 2001; Meredith et al., 2011). Some texts consider the land members of the artiodactyls and the marine members separately due to the great phenotypic divergence that has taken place. Here we consider them together because it more clearly documents an independent lineage's evolution with adaptions to a broad range of environments. The smallest member of this clade is the lesser mouse-deer (body weight of ∼2   kg), and the largest is the blue whale (body weight of ∼115   000   kg), which is also the largest mammal (Matsubayashi, 2003; Small, 1971).

The land artiodactyls include many commonly known animals, but despite this, little research has been done to characterize their neuroanatomy. For the majority of species studied, the external anatomy suggests that there is little emphasis placed on the representation of the hindlimbs and more emphasis on the face. A small cuneate–gracile complex and relatively large trigeminal nuclei have been noted in the giraffe (Badlangana et al., 2007) and sheep (Woudenberg, 1970). Electrophysiological mapping of the neocortex also found a large representation of the mouth and lips in sheep (Johnson et al., 1974). The external anatomy of the face of some species suggests that there may be important tactile regions, such as the oral regions of the hippopotamus, which is studded with prominent tactile hairs, or the large glabrous snout of the pig. Unfortunately, descriptions of the neuroanatomical correlates of these body surfaces are lacking.

During the evolution from a terrestrial ungulate-like mammal to a marine mammal, cetaceans lost functional hindlimbs, greatly elaborated their tail, and transformed their forelimbs from legs into flippers. In addition, some whales and dolphins have specialized hairs and some dolphins have pits on their head region; both of these structures are presumed to be sensory adaptations. In humpback whales (Megaptera novaeangliae), the base of the hair has developed into a visible, knoblike swelling called a tubercle. These tubercles are highly innervated (Yablokov, 1974). Their function may be related to feeling the density of plankton while feeding, detecting low-frequency sound, perceiving water movement, or sensing an yet unknown stimulus (Mercado, 2014). It has been proposed that the pit structures in the Guiana dolphin (Sotalia guianensis) might be electrosensory (Czech-Damal et al., 2012). Another unusual feature in cetaceans is the tusk of narwhals (Monodon monoceros) that is highly innervated by dentinal tubules projecting to the trigeminal nerve. If this is an important sensory apparatus, as some have suggested (Nweeia et al., 2014), it should have a significant representation in the trigeminal somatosensory brain stem.

Very little research has investigated the central nervous system representation of any aspect of the somatosensory system in cetaceans. However, it has been descriptively noted that cuneate–gracile complex is proportionally reduced in size compared to other mammals [this has been noted in a short-beaked common dolphin (Delphinus delphis) (Hatschek and Schlesinger 1902), a bottlenose dolphin (genus: Tursiops) (Wilson, 1933), and a baleen whale (genus: Balaenoptera) (Norris and Sciences, 1966)]. The SpV and the PrV also appear reduced in size compared to other mammals but not as much as the dorsal column nuclei (Wilson, 1933; Hatschekand Schlesinger 1902; Norris and Sciences, 1966). The size of the trigeminal nucleus in the baleen whales may be related to the large size of their head (about one-third of the body) (Norris and Sciences, 1966). No mention of any special representation of the tubercles, pits, or tusk could be found.

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Psychology Consult

Robert J. Maiden , ... Benjamin A. Bensadon , in Psychology and Geriatrics, 2015

Patient Safety

If ranked as a disease, adverse drug reactions would be the fifth leading cause of death in America (Petrone & Katz, 2005, p. 757–758). Iatrogenic impact of potentially inappropriate prescribing is a major threat to patient safety, increasingly a national priority according to an IOM report in 2000, To Err Is Human: Building a Safer Health System (Kohn, Corrigan, & Donaldson, 2000).

Geriatric patients are particularly vulnerable to medication-related error involving polypharmacy and nonadherence. Older adults are the greatest consumers of medication, with more than 94% of women over the age of 65 taking at least one daily prescription medication and 12% taking 10 or more (Kaufman, Kelly, Rosenberg, Anderson, & Mitchell, 2002). Not only do elderly patients consume more medication than their younger counterparts, but aging-associated pharmacokinetic and pharmacodynamic changes increase susceptibility to drug-drug and drug-disease interactions that often are not considered until it is too late (Mallet, Spinewine, & Huang, 2007). Research has found 35% of ambulatory older adults experience adverse drug reactions on a yearly basis and 29% of these require physician evaluation in an emergency room or hospital (Petrone & Katz, 2005).

Geriatricians have identified pharmaceutical risks to guide prescribing clinicians regarding potentially inappropriate medications, particularly for nursing home residents, via the "Beers Criteria" (Beers et al., 1991; Fick et al., 2003). But while this guide may help safeguard against physician prescription-related errors, it cannot sufficiently prevent patient error, such as that caused by misunderstanding directions and subsequent failure to take medications as prescribed, a critical behavioral component of effective medical treatment. The clinical relevance and poor management of patient adherence has been discussed by psychologists for decades (DiMatteo, 2004a), as has the key benefit conferred by social support (DiMatteo, 2004b). Multimorbid older patients are especially vulnerable and may be seriously challenged by complex medication regimens and/or adverse side effects. For some chronic diseases, in which the presence and severity of symptoms may fluctuate or go unnoticed (e.g., hypertension), ensuring appropriate adherence can be extremely difficult for any clinician, especially since standard frequency of ambulatory outpatient contact with primary care physicians is 1–3 visits per year. Some have estimated that only 29–59% of elderly patients are able to take their medications as prescribed (Stewart & Caranasos, 1989). In response, many geriatric physicians (e.g., Rayner, O'Brien, & Schoenbachler, 2006) advocate the "start low and go slow" approach.

In some cases patients may not feel motivated to adhere to their medication regimen or adjust their lifestyle (e.g., diet, exercise). As described elsewhere in this publication, motivational interviewing (MI), a behavioral intervention created by psychologists and most often used by psychologists, has proven effective for negotiating patient ambivalence about behavior change related to health and illness. Rubak and colleagues (2005) conducted a systematic review and meta-analysis of 72 randomized control trials that revealed a significant, clinically relevant effect of MI in 75% of studies reviewed. Impact was comparable on both physiologic and psychological conditions and more likely to be achieved when MI was conducted by physicians and psychologists than by other professionals (80% vs. 46%). Consulting psychologists can conduct such interventions and train other providers to do so.

Harvard surgeon Atul Gawande has described the enormous human and financial costs associated with adhering solely to a medical model of geriatric care (Gawande, 2014). Others have also shown that lack of mental health input leads to higher health care utilization and costs, greater functional impairment, increased utilization of staff time, patient non-adherence, increased mortality, and reduced quality of life (Hyer and Shah, 2009; Ormel et al., 1998; Rodriguez, 2013). A growing number of physician and nonphysician geriatric specialists (e.g., Bensadon & Odenheimer, 2014; George & Whitehouse, 2014; Kathol, deGray, & Rollman, 2014) have therefore recommended psychosocial interventions as appropriate first-line treatment of conditions for which unnecessary medical and surgical services continue to be used at an annual cost approaching 350 billion dollars. Even among the most seriously mentally ill, "two-thirds of the patients receive no care at all, and those who do, receive care in the medical and not the behavioral care sector" (Hyer & Shah, 2009, p. 172).

Kathol and colleagues (2014) propose the following patient-centered LTC model: 1. The focus should be on complex medical illnesses such as diabetes, asthma, heart disease and patients with high health care costs rather than on screening everyone for behavioral problems; 2. Resources should be administered using a fully integrated model; 3. Behavioral health practitioners (i.e., psychologists) should be well trained in evidence-based methods; and 4. The latest technology should be employed, including telecommunications.

Targeting medically complex and high-utilization patients has led to functional and cost improvements, consistent with the triple aim of enhanced patient experience, population health, and lower per capita cost.

Case Example: The Pink Giraffe

A psychologist was consulted to engage a patient who was convinced she was psychotic after seeing a pink giraffe in her hospital room. Physician response was to simply increase her medication. But only after spending time listening to her narrative did it become clear she had already been prescribed too much amitriptyline and had suffered an adverse reaction. With continued discussions, the patient was able to work through her fears and finally accept that her hallucinations were prompted by an allergic reaction to her medicine, which had caused her delirium. Psychologist consultation was ideal for several reasons. Training and experience enabled the psychologist to avoid prematurely judging the patient's report of hallucinations as evidence of psychosis. Instead, the patient was given a chance to share her story with a nonjudgmental doctor well-trained in active listening. The more the patient was reassured and felt understood, the more she trusted, and subsequently the more details she shared about her unsettling symptoms and related fear. This nonpharmacologic intervention, characterized by health literacy information exchange and nonjudgmental empathic listening, successfully broke the all too frequent cycle of harmful and ineffective but ongoing medication administration (see Campanelli, 2012).

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Female Reproduction

Jonathan A. Green , Madison E. Hennessy , in Encyclopedia of Reproduction (Second Edition), 2018

PAGs Are Restricted to the Cetartiodactyla Order

The PAG gene family is restricted to the Cetartiodactyla order. Animals within this order include swine, giraffes, cattle, bison, sheep, whales, and related species. When conducting a phylogenetic analysis of all known PAGs in public databases, there are currently over 250 total PAG genes present. Fig. 1 represents a simplified phylogenetic tree that illustrates the relationship between PAGs and other mammalian aspartic proteinases. Based on current genome sequencing results, PAG genes are most prevalent in ruminant species. In other species of the Cetartiodactyla order the number of PAG genes is smaller. For example, the number of total PAG genes present per species ranges from two genes represented in the genome builds of some whale species, to over thirty known PAG genes in many of the Bovidae (Telugu et al., 2009). It is notable that, when comparing across the different suborders (Suidae, Cetacea, Ruminantia), there is little indication that individual PAGs are conserved across the order. In other words, there are no obvious orthologs across the suborders, and all the PAGs represented by each suborder cluster together in phylogenetic analyses. It is not clear whether that pattern is due to expansion of PAGs in these species after their split from a common ancestor or due to homogenization of PAG genes in these suborders. There is also evidence that certain parts of these proteins (mainly surface loops) are evolving rapidly (Hughes et al., 2000).

The diversity in the number of PAG genes, their lack of orthology and/or their rapid evolution is suggestive that the functional role of PAGs is likely to differ across species within the suborders of the Cetartiodactyla. It is worth noting that all cetartiodactyls possess syn/epitheliochorial placentas that are adaptive forms that are distinct from the ancestral endotheliochorial or hemochorial forms (Wildman et al., 2006). Presumably, the PAGs are performing functions that are important for these unique placental forms. Speculations about those possibilities are elaborated upon below.

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