Lecture 5
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Feedback systems
Overview: Feedback systems involve a disturbance, actuators (effectors), a regulated variable, a sensor, a setpoint, and a feedback loop.
Page 2 components:
Disturbance: external change.
Effector.
Variable.
Sensor.
Feedback loop.
Page 3 details:
(a) Disturbance (example: 17).
(b) Effectors, e.g., sweat glands, thermogenesis, behavior.
Variable (example: 7b).
Setpoint.
Comparator.
Error.
Page 4 flow:
Substrate (c).
Enzyme A.
Error signal.
Enzyme B.
Enzyme C.
Product A.
Product B.
End-product.
Scaling
Page 5: Table 3.1 – Ranges of organism and animal sizes (Mass in grams; Units to scale).
Mycoplasma: $10^{-13}$ g; < 0.1 pg.
Typical bacterium: $10^{-10}$ g; 0.1 ng.
Tetrahymena: $10^{-7}$ g; 0.1 μg.
Amoeba: $10^{-4}$ g; 0.1 mg.
Rotifer: $10^{-4}$ g; 0.1 mg.
Aphid: $10^{-3}$ g; 1 mg.
Bee: $10^{-1}$ g; 100 mg.
Pygmy shrew: $10^{2}$ g; 1 g (note: table shows a size jump toward small mammals).
Hamster: $10^{2}$ g; 100 g.
Human: $10^{5}$ g; 100 kg.
Elephant: $5 imes 10^{6}$ g; 5000 kg (5 tonnes).
Blue whale: $10^{8}$ g; 100,000 kg (100 tonnes).
Page 6: Scaling summary
Many physiological processes vary with organism size.
Not always in a straightforward fashion (e.g., metabolic rate vs locomotion rate do not scale in lock-step with size).
A hamster is roughly 100x more massive than a pygmy shrew, but its metabolic rate is not 100x that of the shrew.
Statement: Metabolic rate tends to be lower as mass increases when correcting for body mass, i.e., mass-specific metabolic rate decreases with size.
Page 7: Other size-related traits
Some traits increase with size and are tightly linked to body size (e.g., bone thickness).
Purpose of scaling: remove body size as a confounding factor to reveal underlying variation (SCALING).
Scaling reveals structural and functional consequences of size changes among otherwise similar organisms.
Page 8–9: Scaling can reveal constraints
Example: Elephants cannot fly because of environmental and physiological constraints.
Concept: Something roughly 20x larger than an elephant could exist in water, where water provides support and aids heat loss.
On land, those constraints create limits on biological possibilities and adaptation.
Page 10: Isometric scaling & Allometry; Log plots
Isometric scaling: a trait varies in direct proportion with body size.
Example: Surface area scales with volume by the exponent ~0.67 (i.e., 2/3).
In general: where is the mass exponent.
Log form: where
= intercept (proportionality constant),
= slope (mass exponent).
Why use log plots? They linearize power-law relationships and help compare scaling across sizes.
Page 11: Real-world scaling patterns
In reality, few properties vary isometrically; most show allometric variation between body size and the trait of interest.
The plotted curve often does not pass through the origin.
Page 12: Scaling and confidence intervals
Confidence intervals are often shown on log plots.
Log plots reduce large differences on linear scales, so small slope differences can be biologically important.
Many previously interpreted adaptive differences may reflect scaling differences.
Organisms deviating from the line (well above or below) are often targets of focused study for insights into constraints and biological possibilities.
Page 13: Metabolic rate and body size (Figure 3.5)
Metabolic rate (MR): rate of calories expended.
Body size is a strong predictor of MR; MR typically follows an allometric relationship (no line through the origin).
About 90% of variation in MR observed within groups is attributable to variation in body mass; taxonomic affiliation also contributes to differences.
Page 14: Mice, small birds, and marmots (Figure 3.5)
Mice and small birds: birds of similar body size have higher MR than mammals of similar size; small birds must feed more frequently or consume energy-dense foods to support high MR.
Marmots: low MR for their size, often at high elevations; spend many months hibernating; store fat to support long periods of inactivity.
Page 15: Sex differences in MR
On average, women have lower MR than men.
When sexes are combined, the data align with the overall MR line, indicating a sex-specific difference in MR.
Page 16: Implications of size on heat and water loss
Larger animals have a lower surface area:volume ratio.
This affects heat loss/retention and evaporative water loss.
Page 17: Oxygen consumption (Figure 3.6)
Oxygen consumption (as a proxy for MR) shows a slope similar to MR vs body mass.
When MR is scaled for body mass, oxygen consumption per unit mass decreases with increasing body mass (specific consumption) as long as the slope is less than 1.
Other traits show similar scaling relationships.
Page 18: Specific cost of locomotion (Figure 3.10)
The cost of locomotion scales with body mass in a way that is broadly consistent across taxa.
Within and among taxonomic groups, there are similar scaling patterns for locomotion costs.
Page 19: Locomotion mode and endothermy/ectothermy differences
The mode of locomotion affects efficiency; some modes are more efficient than others.
Endotherms and ectotherms show differences in scaling of locomotion costs.
Figure 3.14 indicates that while moving a larger body costs more energy, more ground may be covered per unit of movement depending on mode and physiology.
Connections and implications
Practical implications:
Scaling informs comparative physiology, ecology, and evolutionary biology by isolating size effects from other traits.
Understanding MR scaling helps explain energy budgets, feeding strategies, and activity patterns across species.
Conceptual implications:
Distinguishing isometric vs allometric relationships clarifies why simple ruler-based comparisons can be misleading.
Allometry highlights constraints that shape the evolution of form and function.
Ethical/philosophical notes:
When applying scaling concepts to real-world biology, consider limitations of cross-species generalizations and the importance of taxonomic context.
Recognize that deviations from allometric predictions can reveal novel biology requiring deeper investigation.
Key equations and definitions (summary)
Allometric relation (general form):
where is the trait, is body mass, is a constant, and is the mass exponent.Log-transformed linear form:
This yields a straight line with intercept and slope on a log-log plot.Isometric scaling example (surface area vs volume):
Mass-specific metabolic rate concept:
Mass-specific MR typically decreases with increasing body mass (i.e., MR per unit mass decreases as increases).
Oxygen consumption scaling (concept):
If MR scales as with , then specific (per-mass) MR decreases with increasing ; similarly for oxygen consumption with a slope similar to MR.
Conceptual terms to remember:
Isometry: trait changes in direct proportion to body size ().
Allometry: trait changes with body size at a rate different from direct proportion ().
Mass-specific metabolic rate: MR per unit body mass; tends to decrease with larger body mass.