Module 3
模組3
Physiological measures of
animal welfare (1):
動物福利之生理學測定法（一）
Autonomic Nervous System
自主神經系統
This module will enable you to
此模組的教學目標
• Understand the relationship between welfare
and physiology 了解動物福利與生理學間的關係
• Examine how the autonomic system (ANS) is
associated with changes in welfare 檢測當福利改
變時如何牽動自主神經系統的變化
• Understand how measurement of ANS
responses can be used to assess welfare 了解
自主神經系統的反應測試如何應用於評估福利狀態
• Identify some pros and cons of using ANS
measures to assess welfare 找出使用自主神經系統
測量來評估福利狀態的方法之優缺點
Definition of animal welfare
(physical status)
動物福利之定義（生理狀態）
“Animal welfare is the physical and
psychological state of an animal as regards
its attempt to cope with its environment”
“動物福利是動物對環境的生理與心理之調適。”
Professor Donald M Broom
University of Cambridge, UK
英國劍橋大學教授
Donald M Broom
Change in welfare
動物福利改變
‫װ‬
Change in physical & psychological
states
生理及心理狀態改變
‫װ‬
Physiological responses
產生生理反應
Types of response
反應種類
Change in welfare
當動物福利改變時

Activation of CNS
中樞神經系統受刺激活化


Autonomic nervous
responses
Neuroendocrine
responses
自主神經系統反應
神經內分泌系統反應
Autonomic Nervous System (ANS)
自主神經系統
Sympathetic adrenal
medullary system
(SAM)
Parasympathetic
nervous system
(PNS)
交感神經-腎上腺髓質系統
副交感神經系統
SAM process
交感腎上腺系統機制
Stimulus of CNS
刺激中樞神經系統
Adrenal medulla
Heart; metabolism; etc
腎上腺髓質
心臟、新陳代謝等
兒茶酚胺
Catecholamines
Innervation
神經分佈
Circulation
血液循環
Principal SAM effects
交感腎上腺系統之生理反應效果
• Increased cardiac output: 增加心輸出
– Tachycardia 心搏加快
– Increased cardiac muscle contraction 心肌收縮增強
• Increased flow of blood to muscles:
增加流向肌肉的血量
– Peripheral vasoconstriction 周圍血管收縮
– Contraction of the spleen 脾臟收縮
• Increased air intake: 增加氣體吸取
– Increased respiratory rate & bronchiole relaxation
呼吸速率提升與肺部小支氣管放鬆
Parasympathetic Nervous System
副交感神經系統
• Regulates SAM system
調控交感腎上腺系統
• Reduces cardiac output
減少心輸出
– Bradycardia via effect on
sino-atrial node 藉由在竇房結上作
用達到心搏徐緩的效果
Using the ANS to assess welfare
利用自主神經系統評估動物福利
• ANS response = acute measure
自主神經系統反應 ＝ 急性動物福利變化之測量
• Pathological changes = chronic measure
病理變化 ＝ 慢性動物福利變化之測量
Using the ANS to assess welfare
利用自主神經系統評估動物福利
• Direct measures: 直接測量項目
– Heart rate 心跳速率
– Blood pressure 血壓
– Respiratory rate 呼吸頻率
– Catecholamine levels 兒茶酚胺濃度
• Indirect measures: 間接測量項目
– Adrenal habituation 腎上腺適應化
– Adrenal enzymes 腎上腺酵素
Measurement techniques
and their drawbacks
測量技術與其缺點
• Implanted devices 侵入式儀器
– Invasiveness 嵌入身體組織
• Hands-on measurement
– Restriction 需要保定
– Disturbance 動物干擾
手動測量
(i) Heart rate measurement
心跳測量
Method
Invasive
Restrictive Disturbing
方法
侵入性
限制性
Direct
干擾性






Telemetry



Remote



e.g. stethoscope
直接法（如聽診器）
Tethered
e.g. polar monitor
頸鏈（如極性監測）
嵌入性遙測裝置
遠距遙控裝置
Heart rate
心跳速率
Indicates welfare at that point in time
心跳是當前動物福利之指標
Change in welfare
福利有所改變時
 H.R.
 H.R.
Tachycardia
(Active response)
Bradycardia
(Passive response)
心搏加速（主動反應）
心搏徐緩（被動反應）
Heart rate increase
心跳增加的代表意義
• Sheep heart rate increased on exposure to:
綿羊心跳上升的可能情況：
– Strange person (increase of 45 beats per
minute) 遇到陌生人（增加至45跳/分）
– Strange person and dog (increase of 79 bpm)
遇到陌生人帶著狗（增加到79跳/分）
(Baldock & Sibley, 1990)
Heart rate decrease
心跳減緩的代表意義
• Humans—fainting 當人暈厥的時候
• Rodents, when disturbed by threatening
visual cue or sudden noise 囓齒類動物當受
到視覺上威脅訊息或是突然的巨響干擾時心跳會降低
(Hofer, 1970)
Heart arrhythmias
心律不整的代表意義
• Indicator of chronic welfare changes
心律不整為動物福利慢性變化之指標
• Repeated restraint caused tachycardiac
arrhythmias in squirrel monkeys 反覆的保定造成松鼠
猴心搏加快型的心律不整
(Corley et al., 1973)
• Repeated noise and threatening stimuli caused
bradycardiac arrhythmias in rats 不停歇的噪音和脅迫
性刺激造成大鼠心搏徐緩型的心律不整
(Hofer, 1970)
(ii) Blood pressure measurement
血壓測量
Method
Invasive
Restrictive
Disturbing
方法
侵入性
限制性
干擾性
Cuff: 血壓袖帶（可
綁在囓齒類的尾巴、
耳朵或手臂上）



Catheterisation:
動脈內導管



Implanted Telemetry
遙測裝置


+/-
Blood pressure
血壓
Measure of chronic welfare change
血壓為動物福利慢性變化之測量方法
• Change in welfare alters blood pressure
福利的變化會影響血壓
–Aggression increased blood pressure in mice 攻擊行
為會增加小鼠的血壓
(Henry et al., 1975)
–Daily immobilisation increased blood pressure in
rats 每日限制活動會升高大鼠的血壓
(Lamprecht et al., 1973)
(iii) Respiratory rate
呼吸頻率
• Assessment of present state 評估目前狀態
• Easy to observe 觀察方便
• Closely correlated with heart rate 與心跳有
密切關係
• Lamb respiratory rate  following:
小羊在下列狀況呼吸速度會加快：
– Tail docking 截尾
– Castration 去勢
(Mellor & Murray, 1989)
(iv) Catecholamine measurement
兒茶酚胺濃度測量
• Rapid sampling required 需要迅速採樣
Method
方法
Invasive
侵入性
Disturbing
干擾性
Restrictive
限制性
Variable
變動性
Catheterisation
置入血管內導管




Urine
尿液採樣




Autopsy
屍體解剖




Analysis: High performance liquid chromatography
(HPLC) 分析法：高壓液相層析法
Catecholamines
兒茶酚胺類
• The adrenal medulla releases:
腎上腺髓質分泌的特性：
– Adrenaline is associate with psychological
stimuli 腎上腺素的分泌與生理刺激有關
– Noradrenaline is associated with physical
stimuli 正腎上腺素的分泌也與生理刺激有關
• Very acute measures
需要快速測量
– Immediate release (Rat: 1-2 seconds) 釋放速度
快（大鼠只需要1-2秒）
– Short half-life (Rat: 70 seconds) 半衰期短（大鼠只
有70秒）
Catecholamine examples
舉例說明引發分泌兒茶酚胺的原因
• Rat catecholamine levels  with:
大鼠血中之兒茶酚胺會在下列狀況中升高：
– Cage door opening 飼籠被打開
– Handling and moving between cages 手動換籠
– Restraint (40 fold ) 保定
(Kvetnansky et al., 1978)
• Defeated male guinea pigs show significantly
higher catecholamine levels compared to
victors 雄天竺鼠敗方體內之兒茶酚胺濃度較勝方顯著
升高
(Sachser & Lick, 1989)
Adrenal habituation
腎上腺適應化檢查
• Chronic welfare assessment 慢性的動物福利評
估
• Repeated problem leads to:
重複實驗會造成的問題：
–  catecholamine to that stimulus (habituation) 兒茶酚胺
對相同之重覆刺激反應濃度降低（適應化）
–  catecholamine to other stimuli (sensitisation) 兒茶酚
胺濃度對不同之其他刺激反應濃度增加（敏感化）
• Assessment: 評估方式
– Assess levels continuously 連續重覆刺激的濃度反應
– Before and after stressful period 緊迫期間前後之濃度比較
(註：若未發生適應化，表示出現福利不良之問題。)
Adrenal habituation: example
舉例說明腎上腺適應化現象
• Repeated restraint of rats 大鼠重複保定試驗
• Day 27 compared to day 1 第27天之結果與第
1天的比較
– Restraint induced  catecholamine
production 保定誘發兒茶酚胺濃度減少
– Different stressor induced  catecholamine
production 不同的壓力源卻引發兒茶酚胺增加濃度
(Konarska et al., 1989)
Limitations of ANS measures
自主神經系統測量法所受到的限制
• Other factors causing changes
其他改變生理狀態的因子
– Activity 動物的活動性
– Metabolism 代謝率
– Timing of sample 採樣時間
• Disturbance caused by measurement
採樣時出現的干擾
– Human presence, handling, restraint and sampling
method 其他人員在旁邊、碰觸動物、保定與採樣方法等，都會造成干擾
• Measurement error
測量方法本身之誤差(要多做幾次)
Limitations of ANS measures
自主神經系統測量法所受到的限制（續）
•
Individual differences
-
個體之差異性
High and low responders 體質反應之高低
 Rats (Livezey et al., 1985)
- Dominance status 社會位階之強勢/弱勢狀態
 Tree shrews 尖鼠實驗(von Holst, 1986)
- Sex 性別本身之差異性
 Rats 大鼠實驗(Livezey et al., 1985)
Conclusions
結論
• ANS response = Acute measure
自主神經系統反應 ＝ 急性測量
• Chronic change = Chronic measure
慢性變化 ＝ 慢性測量
• Potential parameters:
潛力參數
– Direct: Heart rate, blood pressure, respiratory rate and
catecholamines 直接參數：心跳、血壓、呼吸頻率、與兒茶酚胺濃度
– Indirect: Adrenal habituation, enzymes & metabolites 間接
參數：腎上腺適應性、酵素與兒茶酚胺之代謝產物
• ANS measures are limited, so care is required in
their assessment 自主神經系統的反應測量是有先天性條件限制
的，在測量時要特別小心注意
Further Reading
進階閱讀
• BALDOCK, N.M. & SIBLEY, R.M. 1990 Effects of handling and
transportation on heart rate and behaviour in sheep. Applied Animal
Behaviour Science 28, 15-39
• CORLEY, K.C., SHIEL, F.O’M., MAUCK, H.P., GREENHOOT, J.
1973 Electrocardiographic and cardiac morphological changes
associated with environmental stress in squirrel monkeys.
Psychsom. Medicine 35, 361-364
• HEMSWORTH PH 2003: Human-animal interactions in livestock
production. Applied Animal Behaviour Science, 81: 185-198
• HENRY, J.P., STEPHENS, P.M. & SANTUSTEBAN, G.A., 1975 A
model of psychosocial hypertension showing reversibility and
progression of cardiovascular complications. Circ. Res. 36, 156-164.
Further Reading
進階閱讀
• HOFER, M. A., 1970 Cardiac and respiratory function during
prolonged immobility in wild rodents. Psychsom. Medicine 32,
633-647
• KONARSKA, M., STEWART, R. E. & McCARTHY, R., 1989
Habituation of sympathetic-adrenal medullary responses
following exposure to chronic intermittent stress. Physiology
and Behavior 45, 255-261
• KVETNANSKY, R., SUN, C. L., LAKE, C. R., THOA, N.,
TORDA, T. & KOPLIN, I. J., 1978 Effect of handling and forced
immobilisation on rat plasma levels of epinephrine,
norepinephrine and dopamine-b-hydroxylase. Endocrinology
103, pp. 1868-1874
• LAMPRECHT, F., WILLIAMS, R.B. & KOPIN, I.J. 1973 Serum
dopamine-beta-hydroxylase during development of
immobilisation-induced hypertension. Endocrinology 92, 953956
Further Reading
進階閱讀
• LIVEZEY, G.T., MILLER, J.M. & VOGEL, W.H., 1985 Plasma
norepinephrine, epinephrine and corticosterone stress
responses to restraint in individual male and female rats, and
their correlations. Neurosci. Lett. 62, 51-56
• MELLOR, D.J. & MURRAY, L., 1989 Effects of tail docking and
castration on behaviour and plasma cortisol concentrations in
young lambs. Res. Vet. Sci. 46, 387-391
• SACHSER, N. & LICK, C. 1989 Social stress in guinea pigs.
Physiology and Behavior 46, 137-144
• VON HOLST, D., 1986 Vegetative and somatic components of
tree shrews’ behaviour. J. Auton. Nerv. Syst.; Suppl.: 657-670