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大學部 生態學與保育生物學學程 (必選) 2010 年 秋冬 生物時鐘(Biological clocks) ─動物行為學 (Ethology) 鄭先祐(Ayo) 國立 臺南大學 環境與生態學院 生態科學與技術學系 教授 Ayo NUTN Web: http://myweb.nutn.edu.tw/~hycheng/ Part 1. 動物行為的研究途徑 (個體行為) 歷史背景 (History of the Study of Animal Behavior ). 基因分析 (Genetic Analysis of Behavior ). 天擇 (Natural Selection and Behavior ). 學習與認知 (Learning and Cognition.) 生理分析 (Physiological Analysis) (一) 神經細胞 (Nerve Cells and Behavior ). (二) 內分泌系統 (The Endocrine System). 發育(The Development of Behavior ). Ayo 教材 (動物行為學 2010) 2 Part 2. 存活 (與環境的互動關係) 生物時鐘 (Biological Clocks) 導航機制 (Mechanisms of Orientation and Navigation) 空間分佈的生態學與演化學 (The Ecology and Evolution of Spatial Distribution) 覓食行為 (Foraging Behavior) 抗掠食行為 (Antipredator Behavior) Ayo 教材 (動物行為學 2010) 3 08 生物時鐘 (Biological clocks) Clock-controlled rhythms Rhythmic behavior The clock versus the hands of the clock Advantages of clock-controlled behavior Adaptiveness of biological clocks Organization of circadian systems Human implications of circadian rhythms Ayo 教材 (動物行為學 2010) 4 Animals have internal clocks Hamsters (倉鼠), as well as all other animals, have an internal, living clock Its bouts of activity alternate with rest It is so regular that it is described as an activity rhythm Ayo 教材 (動物行為學 2010) 5 Most animals can measure time Biological clocks have been found in every eukaryotic organism tested As well as in cyanobacteria The rhythmical nature makes sense in the light of evolutionary principles Ecological conditions vary tremendously at different times of the cycle It is adaptive to predict upcoming changes in a cycle (i.e. upcoming darkness or winter), rather than just respond to these events Ayo 教材 (動物行為學 2010) 6 Environmental modifications are extreme but predictable Biological clocks have evolved as adaptations to environmental cycles Biological clocks also provide a mechanism to synchronize various internal processes with other internal processes Rhythms have piqued(引發) the interest of scientists studying Their adaptive value and evolution Genetic underpinnings (基因基礎) Hormonal control Neural control Ayo 教材 (動物行為學 2010) 7 Properties of clock-controlled rhythms Biological clocks measure time at the same rate under nearly all conditions They can be reset to remain synchronized with environmental cycles In clock-controlled rhythms, cycles continue in the absence of environmental cues (i.e. light-dark and temperature cycles) Instead, the ability to keep time without external cues is due to an internal (endogenous) biological clock Ayo 教材 (動物行為學 2010) 8 Biological rhythms The period (the interval between two identical points in the cycle) of the rhythm can become longer or shorter Circadian rhythm: a daily rhythm (24 hours in nature) Circalunidian: a lunar day (tidal) rhythm Circamonthly: a monthly rhythm Circannual: an annual rhythm Free-running period: a circadian period length under constant conditions It is not manipulated by environmental cycles Ayo 教材 (動物行為學 2010) 9 Entrainment to environmental cycles Biological rhythms are not exactly as long as natural cycles A light-dark cycle: the most powerful phase-setting agent in circadian rhythms Nonphotic cues (i.e. social interactions, feeding schedules, or temperature cycles) also play a part Manipulating the light-dark cycle resets (entrain) the biological clock A brief light pulse during early circadian night resets the clock In nature, the clock is reset by light at dawn and dusk each day Ayo 教材 (動物行為學 2010) 10 Light sampling behavior resets the circadian clock Flying squirrels are nocturnal Their circadian clock wakes them at twilight (薄暮) If it is still light outside, the squirrel returns to its nest to sleep and its circadian clock is reset Activity begins slightly later the next night The squirrel’s activity rhythm entrains to the light- dark cycle with only a few minutes of light exposure This phase adjustment also causes the squirrel’s onset of activity to follow sundown as it changes through the year Ayo 教材 (動物行為學 2010) 11 Biological clocks are insensitive to temperature changes If biological clocks were affected by changes in temperature: They would run at different rates at different times of day Cave dwelling, insectivorous bats of the temperate zone roost in cool, deep caves during the day Their body temperature drops while resting to conserve metabolic energy Their temperature rises when they leave the cave to forage The biological clock remains accurate Ayo 教材 (動物行為學 2010) 12 Daily rhythms Nocturnal animals are busiest at night Hamsters, cockroaches, bats, mice, and rats Diurnal animals are active during the day Most songbirds and humans Crepuscular animals are active primarily at dawn and dusk Ayo 教材 (動物行為學 2010) 13 Lunar day rhythms As the moon passes over the surface of the earth, its gravitational field draws up a bulge in the ocean waters Causing high tides when they reach the shoreline There are usually two high tides each lunar day one every 12.4 hours The tides cause some dramatic changes in the environment, Particularly for organisms living on the seashore Rhythms synchronized with tides are lunar day rhythms Ayo 教材 (動物行為學 2010) 14 Fiddler crab activity is synchronized with tidal changes Fiddler crabs are active during low tide In search of food and mates Before high tide, the crabs return to their burrows A fiddler crab’s behavior in the laboratory remains rhythmic Periods of activity alternate with quiescence every 12.4 hours The usual interval between high tides Ayo 教材 (動物行為學 2010) 15 An activity rhythm of a fiddler crab (Uca pugnax). Althouth the crab was maintained in constant darkness and temperature(20℃). Ayo 教材 (動物行為學 2010) 16 Semilunar rhythms The gravitational field of the sun also influences the height of the tides The highest tides are caused when the gravitational fields of the moon and the sun operate together At new moon and full moon, the earth, the moon, and the sun are in line The gravitational fields of the sun and the moon augment each other Ayo 教材 (動物行為學 2010) 17 The effect of the Ayo 教材 (動物行為學 2010) relative positions of the earth, moon, and sun on the amplitude of tidal exchange. (a) at the times of new and full moons, (b) during the first and last quarters of the 18 Spring and neap tides Spring tides: the highest high tides and lowest low tides at new and full moons Neap tides: periods of lowest high tides and highest low tides at the quarters of the moon Some organisms possess a biological clock that helps them predict the times of spring or neap tides In the tiny chironomid midge, emergence of adults from their pupal cases is programmed to coincide with tidal changes. Ayo 教材 (動物行為學 2010) 19 chironomid midge Chironomidae (informally known as chironomids or nonbiting midges) are a family of nematoceran flies with a global distribution. Many species superficially resemble mosquitoes but they lack the wing scales and elongate mouthparts of the Culicidae. This is a large group of insects with over 5000 described species and 700 species in North America alone. Ayo 教材 (動物行為學 2010) 20 Monthly rhythms A synodic lunar month: the interval from full moon to full moon (29.5 days) The time it takes the moon to revolve once around the earth Some organisms can program their activities to occur at specific times during this cycle The ant lion shows a monthly rhythm in the size of the pit it builds Small arthropods, such as an ant, slide into the pit At full moon, it constructs larger pits This is a clock-controlled rhythm - not a response to an environmental factor Ayo 教材 (動物行為學 2010) 21 Pit size of ant lion A monthly Ayo 教材 (動物行為學 2010) rhythm in the pit size of the predatory ant lion. Monthly rhythms in the pit size of 50 ant lions maintained in constant condition in the laboratory. 22 Annual rhythms Annual biological clocks are important In timing migration To prepare animals physiologically for migration and reproduction A garden warbler in the laboratory with constant temperature and unvarying day length (12 hours light 12 dark) lacks obvious seasonal cues During summer and winter months, it limits activity to daylight hours It becomes active at night during autumn and spring, when it would normally be migrating Ayo 教材 (動物行為學 2010) 23 Annual cycles in migratory restlessness, body weight, testis size, and molting in a garden warbler held in constant lightdark cycle and at a constant temperature. Ayo 教材 (動物行為學 2010) 24 Caged birds have zugunruhe Zugunruhe (migratory restlessness): a cage-adapted nocturnal form of migratory activity The timing function is very important for birds wintering close to the equator A constant photoperiod and variable rainfall and food abundance don’t provide proper cues to signal the time to begin migrating Ayo 教材 (動物行為學 2010) 25 The annual clock affects physiology It readies birds for migration and reproduction A bird gets fatter during winter to provide fuel for the spring migration It molts during winter Its testes enlarge for summer reproductive activity These cycles are free-running for many years in constant conditions The length of the cycle is slightly longer or shorter than a year Ayo 教材 (動物行為學 2010) 26 Circadian vs. annual clocks Seasonal changes may be controlled by a response to the changing photoperiod Shortening days during winter months and increasing daylight of spring and summer Measuring a change in daylength requires only a circadian clock A rhythm controlled by an annual clock continues to be rhythmic even without changing day length Ayo 教材 (動物行為學 2010) 27 Stop and think When one wants to determine whether a daily, tidal, or lunar rhythm is controlled by an endogenous clock, the organism is placed in constant light or constant darkness When one wants to determine whether an annual rhythm is controlled by an annual clock, the animal is kept in a constant photoperiod Why are the procedures different? Ayo 教材 (動物行為學 2010) 28 Anticipation of environmental change Why time an event with a biological clock rather than responding directly to periodic environmental fluctuations? To anticipate periodic environmental changes To synchronize behavior with other events To measure an interval of time Ayo 教材 (動物行為學 2010) 29 Anticipation of environmental change An animal can anticipate change and prepare for it Adult fruit flies emerge from their pupal cases at dawn, when it is cool and moist They can expand their wings with a minimal loss of water If they waited until later, water loss to the arid air could prevent the wings from expanding properly Ayo 教材 (動物行為學 2010) 30 Synchronization of behavior Allowing a clock to control an event allows behavior to be synchronized with a factor in the environment that cannot be sensed directly Bees time their flights to visit flowers that are open(開花) only during certain times A bee can not use vision or olfaction to determine whether flowers far from the hive are open Ayo 教材 (動物行為學 2010) 31 Bees visit open flowers After training, bees visited a feeding station only during learned hours With their biological clock, bees time their visits to flowers so they arrive when the flower is secreting nectar And gather the maximum amount of food with the minimum effort Ayo 教材 (動物行為學 2010) 32 (b) bees were trained to come to a feeding dish only at the specific times at which food was made available. (c) after six days of training, the feeding dishes were left empty and the number of bees arriving throughout the day was recorded. The bees arrived at the feeding station only when food had been previously present. Ayo 教材 (動物行為學 2010) 33 Continuous measurement of time Measuring passage of time is crucial to time-compensated orientation A honeybee’s dance indicates the direction to a nectar source Telling other bees the flight bearing relative to the sun Because the sun is a moving reference point, the honeybee’s biological clock provides the information for The time of day when it discovered the nectar How much time has passed since then Ayo 教材 (動物行為學 2010) 34 Organization of circadian systems A complex nervous system or endocrine system is not an essential component of the biological clock Single cells, unicellular organisms and cells that make up tissues and organs have their own independent clocks There is no such thing as “the” biological clock Clocks are scattered throughout an animal’s body Fruit flies have a multitude of independent clocks throughout their bodies They respond to changes in light-dark cycles without any help from the head Ayo 教材 (動物行為學 2010) 35 Fruit flies have multiple clocks Glow rhythms indicate per gene activity synchronizes with light-dark cycles And continue in constant darkness with a free-running period length Separate cultures of body parts exposed to the same light-dark cycle glow in unison(一致、合諧) Each piece of cultured tissue has its own independent clocks and these clocks have their own photoreceptors Ayo 教材 (動物行為學 2010) 36 The insect brain is not needed as a master clock To synchronize rhythms throughout the body. Clocks in different cells run at slightly different rates without a light-dark cycle Independent clocks gradually become asynchronous When exposed to a new light-dark cycle, the clocks throughout the fly entrain within one cycle and the glow becomes rhythmic again In nature, asynchrony among peripheral clocks is not a problem Fruit flies have an environmental light cycle that can synchronize independent clocks Because each has its own photoreceptor Ayo 教材 (動物行為學 2010) 37 Animals have at least one master clock Most rhythmic animals have multiple independent peripheral clocks throughout the body How are an individual’s clocks synchronized so all rhythmic processes occur at the appropriate time Relative to one another and the environment’s cycles? At least one “master” clock in the brain is entrained by the light-dark cycle It regulates other clocks through the nervous and/or endocrine system Ayo 教材 (動物行為學 2010) 38 The master clock: the general scheme The general scheme of circadian organization One clock or several interacting clocks function as master clocks to synchronize peripheral clocks The output from the master clock(s) can be neural or hormonal The clocks are set to the right time because photoreceptors convey information on the light-dark cycle to the clock(s) Peripheral clocks generate the rhythmic output, which may feedback on and affect the master clock(s) Ayo 教材 (動物行為學 2010) 39 The master clock: photoreceptors What photoreceptors are responsible for entrainment? Mammalian eyes contain photoreceptors for light entrainment These photoreceptors are in a different part of the retina than those involved in vision Information about lighting conditions reaches the clock through the retinohypothalamic tract (RHT) A bundle of nerve fibers connecting the retina with the hypothalamus Ayo 教材 (動物行為學 2010) 40 The master clock: where is it? The circadian system in mammals is a hierarchy of clocks The SCN (suprachiasmatic nucleus) is the master biological clock Evidence that the SCN is the master clock It is independent: SCN activity remains rhythmic in cultures Spontaneous electrical firing of individual neurons is rhythmic in constant conditions, each of them with a slightly different period The SCN is a self-sustaining oscillator that instills rhythmicity in other brain regions through neural connections Ayo 教材 (動物行為學 2010) 41 The master biological clock of mammals SCN The suprachiasmatic nucleus or nuclei (SCN), is a tiny region on the brain's midline, situated directly above the optic chiasm. It is responsible for controlling circadian rhythms. The neuronal and hormonal activities it generates regulate many different body functions in a 24-hour cycle, using around 20,000 neurons. The SCN, pine cone shaped and the size of a grain of rice, interacts with many other regions of the brain. It contains several cell types and several different peptides (including vasopressin and vasoactive intestinal peptide) and neurotransmitters. Ayo 教材 (動物行為學 2010) 42 The suprachiasmatic nucleus (SCN) in the brain (a) the firing rate of a single neuron from the SCN continues fo fire rhythmically in tissue culture. (b) photoreceptors in the mammalian circadian system reach the SCN via the retinohypothalmic tract (RHT). Ayo 教材 (動物行為學 2010) 43 Nerve activity in specific brain regions (the caudate and the suprachiasmatic nuclei) of a rat before and after isolating an “island” of brain tissue containing the SCN. The SCN instills rhythmicity Ayo 教材 (動物行為學 2010) 44 The master clock regulates other clocks When an SCN from a conspecific is transplanted into rats or hamsters that have been made arrhythmic by destroying their own SCN Their activity becomes rhythmic once again The period length of the restored activity rhythm matches that of the transplanted SCN The SCN is the clock that provides timing information And not just a component needed to make the host’s clock function Ayo 教材 (動物行為學 2010) 45 The genetic basis of circadian timing What are the molecular gears that make the clock tick? Rhythmic gene activity is involved in the clock mechanism The products of one gene or set of genes activate or inhibit the activity of other genes Which in turn affect the activity of the first genes Creating a self-regulated feedback loop of gene activity measuring approximately 24 hours Ayo 教材 (動物行為學 2010) 46 The genetic basis of the circadian cycle Two proteins—Clock and BMAL1—bind together, forming a complex that enters the nucleus The Clock/BMAL1- complex turns on the activity of the period (per) and the cryptochrome (cry) genes The protein products of these genes (Per and Cry) bind with the protein product of the tau gene to form a complex Ayo 教材 (動物行為學 2010) 47 The genetic basis of circadian timing in mammals consists of two feedback loops in gene activity. Ayo 教材 (動物行為學 2010) 48 A 24-hour feedback mechanism The Per/Cry/Tau complexes suppresses action of the Clock/ BMAL1 complex Resulting in less activity of per and cry With less Per and Cry produced and the degradation of Per, Cry and Tau The level of the Per/Cry/Tau complex declines With less inhibition of their activity, per and cry are turned on again This cycle takes about 24 hours to complete Ayo 教材 (動物行為學 2010) 49 Peripheral clocks The SCN may be the master biological clock But other circadian clocks throughout the body keep their own internal time Rhythms persist in cell cultures from the SCN, liver, and lung And in fibroblasts (generic cells) in cultures Perhaps all cells have personal clocks Ayo 教材 (動物行為學 2010) 50 A model of circadian organization in mammals. Ayo 教材 (動物行為學 2010) 51 Clock output The SCN entrain peripheral circadian oscillators So they are correctly set to environmental time The phase relationship between rhythmic output of the SCN and rhythmic clock genes in peripheral tissues varies Peak clock gene expression occurs at distinct times of the day and varies in different tissues The clock may not directly cause the rhythmic output of peripheral tissues Most neural connections are to the hypothalamus and autonomic nervous system, which influences hormone levels Ayo 教材 (動物行為學 2010) 52 Signals from the SCN can be neural The SCN has at least two neural output pathways that affect rhythms One pathway goes to the preoptic nucleus of the hypothalamus Controls the rhythm in ovulation, but not the activity rhythm The second pathway leads to the paraventricular nucleus in the hypothalamus Integrates neuroendocrine and autonomic functions And then leads to the pineal, which produces melatonin Ayo 教材 (動物行為學 2010) 53 Melatonin (退黑激素) 退黑激素(Melatonin)有人叫聚黑激素,它是腦部「松 果體」所分泌的一種激素。它可使青蛙皮膚色素細 胞內之黑色素顆粒聚合於細胞核附近(故稱為聚黑 激素),因而使皮膚顏色看起來較淡。 光線經過視網膜神經細胞再傳至下視丘,再經交感 神經而傳至松果體,抑制退黑激素的分泌。反之, 黑暗則可促使退黑激素的分泌。 下視丘內之一些細胞有如「生物時鐘」般使松果體 之退黑激素分泌出現晝夜韻律之差異,一般晚上入 睡後其血中濃度為白天的六倍。退黑激素在血中的 半衰期甚短,約為半分鐘至5分鐘之間,主要在肝 臟內代謝而其代謝物則由尿液排出。 Ayo 教材 (動物行為學 2010) 54 The influence of light and darkness on circadian rhythms and related physiology and behavior through the SCN in humans Ayo 教材 (動物行為學 2010) 55 Signals from the SCN can be hormonal Reproductive responses to the length of day and rhythms in hormones require neural connections And also small molecules that diffuse to their target Activity rhythms are caused by the SCN’s release of chemical signals to other parts of the brain without neural connections In mammals, the rhythmic production of two hormones (melatonin and glucocorticoids) entrain peripheral oscillators Ayo 教材 (動物行為學 2010) 56 糖皮質激素(Glucocorticoid) 糖皮質激素(Glucocorticoid),學名叫做「腎上 腺皮質素」,由於可用於一般的抗生素或消炎藥 所不及的病症,如SARS、敗血症……等,是由 腎上腺皮質分泌的一類甾體激素,具有調節糖、 脂肪、和蛋白質的生物合成和代謝的作用,還具 有抗炎作用,稱其為「糖皮質激素」是因為其調 節糖類代謝的活性最早為人們所認識。 Ayo 教材 (動物行為學 2010) 57 Hormones entrain peripheral oscillators Neural connections from the SCN cause the pineal to produce melatonin Melatonin amplifies the body temperature rhythm, facilitates sleep, and controls photoperiodic responses Neural connections from the SCN to the anterior hypothalamus initiate hormonal events Glucocorticoids were produced by the adrenal gland Glucocorticoids: steroid hormones that control many physiological functions Ayo 教材 (動物行為學 2010) 58 Stop and think Light cannot reset the clock in the SCN of people who are totally blind Consequently, their sleep-wakefulness rhythms drift out of phase with the day-night cycle They are often sleepy during the day or wide awake at night In one experiment, blind people were able to set their clocks by taking a dose (10 mg) of melatonin at bedtime Why do you think this is possible? Ayo 教材 (動物行為學 2010) 59 Human circadian rhythms: jet lag Jet lag: the most familiar way circadian clocks affect humans A syndrome of effects that includes decreased mental alertness and increased gastric distress Caused by a disruption of circadian timing After traveling across time zones, your biological clocks are still set to the local time of your home The clocks gradually adjust to the day-night cycle in the new locale Ayo 教材 (動物行為學 2010) 60 Resetting biological clocks takes time After traveling, it may take several days to reset your biological clock The more time zones crossed, the longer it takes to reset the clock Not all body functions adjust at the same rate Phase relationships in physiological processes are upset Your body time is out of phase with local time Your rhythms may peak at inappropriate times You suffer psychological and physiological disturbances Ayo 教材 (動物行為學 2010) 61 Stop and think If you were traveling from Tampa Florida to San Diego California to compete in an important athletic event, what steps could you take before you left to minimize jet lag? If you could choose the time of the event, would you choose morning or afternoon? Why? Ayo 教材 (動物行為學 2010) 62 Human circadian rhythms: human health Nearly every physiological process in humans is rhythmic Each process peaks at the appropriate time of day Certain acute medical conditions occur at a certain time of day Most heart attacks and strokes occur between 6 AM and noon Blood pressure rises, platelets become stickier and more likely to form blood clots, and the mechanism that breaks down blood clots is least active Ayo 教材 (動物行為學 2010) 63 Human circadian rhythms: human health Asthma (氣喘) attacks mostly occur at night Levels of epinephrine (a hormone that causes the air tubules to dilate), and cortisol (a hormone that suppresses the immune system) are low The hormone leptin decreases appetite and ghrelin increases appetite Sleep-deprived humans have low leptin levels and high ghelin levels, giving them a heartier appetite than usual Ayo 教材 (動物行為學 2010) 64 Circadian rhythms, sleep and energy The clock gene is part of the circadian mechanism in mammals Clock mutant mice eat more than normal mice They gained weight, developed high cholesterol, blood sugar and triglycerides, low insulin, and bloated fat cells Ayo 教材 (動物行為學 2010) 65 New possibilities for treating diseases Discovering the link between the circadian clock and metabolism has opened new possibilities for treating diabetes and obesity The activity of a gene called SIRT1 is clock controlled It is modulated by how many nutrients a cell is consuming SIRT1 responds to the energy state of a cell and transmits that information to the clock by binding to the BMAL1-Clock complex Helping to explain why lack of sleep can increase hunger and lead to obesity and diabetes Ayo 教材 (動物行為學 2010) 66 Summary Clocks evolved as adaptations to environmental cycles Periods are called circadian, circalunidian, or circannual The free-running period: the period length in constant conditions Entrainment adjusts the period length and the rhythm’s phase Daily rhythms can be entrained to light-dark and temperature cycles Rhythmic processes match geophysical periods and are caused by an internal biological clock Processes become rhythmic when coupled to the biological clock Ayo 教材 (動物行為學 2010) 67 Summary A biological clock measures time for: (1) anticipation of the environmental changes (2) synchronizing the behavior with unsensed events (3) continuous measurement of time A functional clock enhances survival One or more master clocks (SCN) regulates other, slave clocks The master clock regulates activity through nerves and chemicals The genetic basis of circadian clocks involves feedback loops The SCN controls the pattern of gene expression in tissues The human circadian clock is related to health in several ways Ayo 教材 (動物行為學 2010) 68 問題與討論 [email protected] Ayo 台南 NUTN 站 http://myweb.nutn.edu.tw/~hycheng/ Ayo 教材 (動物行為學 2010) 69