Table of Contents
Overview
Behavioral biology (ethology) investigates how and why animals, including humans, behave the way they do. This chapter outlines the main goals of this field and the basic methods used to record, analyze, and interpret behavior. Later chapters will address particular types of behavior and their development; here we concentrate on what behavioral biologists aim to find out in general, and how they study it.
Goals of Behavioral Biology
Behavioral biology pursues several interconnected goals. These can be grouped into four broad questions that can be asked about any behavior:
- What causes this behavior in the short term?
- How does the behavior develop over the lifetime of an individual?
- What is the function or survival value of the behavior?
- How did this behavior evolve in the species?
These are often called the “four levels of explanation” for behavior.
Proximate Goals: Immediate Causes and Mechanisms
Proximate (immediate) explanations focus on how a behavior is produced here and now.
1. Identifying triggers and stimuli
A central goal is to discover which environmental stimuli or internal states trigger a particular behavior:
- External triggers: light, sounds, odors, movements, presence of conspecifics (members of the same species), predators, prey, etc.
- Internal states: hunger, hormone levels, fatigue, reproductive status.
Behavioral biologists want to know:
- Which specific features of a stimulus are crucial (for example, color vs. shape)?
- How strong a stimulus must be to elicit a response (thresholds)?
- How animals distinguish between relevant and irrelevant stimuli.
2. Understanding physiological and neural mechanisms
Another goal is to connect behavior to its physical basis in the organism:
- Which sensory organs detect the relevant stimuli?
- How is information processed in the nervous system?
- How do hormones and other chemical signals modulate behavior?
This includes, for example:
- Relating courtship behavior to hormone cycles.
- Linking escape responses to specific neural circuits.
While detailed mechanisms are explored more deeply in other chapters (e.g., nervous systems, hormones), behavioral biology aims to connect those mechanisms to observable actions.
Ontogenetic Goals: Development of Behavior
Ontogenetic (developmental) explanations ask how behavior changes over an individual’s life:
- Which aspects of behavior are present at birth or hatching?
- Which are acquired through experience (learning, imprinting, practice)?
- How do critical periods (sensitive time windows early in life) shape behavior?
- How do age, growth, and maturation affect behavioral patterns (e.g., juvenile play vs. adult territoriality)?
Behavioral biologists want to chart the “behavioral biography” of an individual:
- When does a behavior appear for the first time?
- How is it refined or modified with experience?
- How stable is it over the lifetime?
Functional Goals: Survival and Reproductive Value
Functional (adaptive) explanations address what the behavior is “good for” from an evolutionary perspective:
- How does a behavior increase an individual’s chances of survival?
- How does it contribute to reproductive success (producing offspring that survive and reproduce)?
- What are the costs and benefits of a behavior (e.g., energy use, risk of predation, missed opportunities)?
Behavioral biologists analyze:
- Feeding strategies: efficiency vs. risk.
- Anti-predator behaviors: camouflage, fleeing, mobbing.
- Social behaviors: cooperation, competition, parental care.
- Mating behaviors: mate choice, courtship displays.
Importantly, the goal is not to assume every behavior is perfectly adapted, but to test whether and how a behavior can be understood in terms of costs and benefits in a given environment.
Phylogenetic Goals: Evolutionary History of Behavior
Phylogenetic explanations ask how a behavior originated and changed over evolutionary time:
- Do closely related species share similar behaviors?
- Are there patterns of behavior that can be traced back to a common ancestor?
- How have changes in environment or way of life shaped behavioral evolution?
Behavioral biologists:
- Compare behaviors across species (comparative method).
- Map behaviors onto evolutionary trees (phylogenies).
- Distinguish between:
- Homologous behaviors: similar due to shared ancestry.
- Analogous behaviors: similar due to similar selective pressures, but independent origins.
This helps reveal whether similar-looking behaviors in different species have the same underlying origin or evolved separately in response to similar challenges.
General Strategies in Behavioral Research
To reach these goals, behavioral biology uses a range of methods. These methods must:
- Record behavior reliably and objectively.
- Allow comparisons between individuals, groups, or species.
- Make it possible to test explanations, not just describe patterns.
Two broad approaches are used:
- Descriptive (observational) methods: document behaviors as they naturally occur, often in the field.
- Experimental methods: manipulate conditions (stimuli, environment, social context) to test specific hypotheses.
Both are essential and are often combined within a single research program.
Observational Methods
Observational methods focus on watching and recording behavior without (or with minimal) intervention.
Ethograms: Catalogues of Behavior
A basic tool is the ethogram, a systematic list of all behavior patterns shown by a species in a particular context.
An ethogram typically includes:
- A clear name for each behavior (e.g., “grooming,” “threat display”).
- A precise description of what it looks like (posture, movements, facial expressions).
- Possibly sketches or standardized symbols.
- Notes on when and where the behavior occurs (context).
Goals of using ethograms:
- Ensure that different observers identify behaviors in the same way.
- Provide a basis for quantitative recording (how often, how long, in what sequence).
- Allow comparisons across individuals, populations, and species.
Sampling Methods: Who or What to Observe
Since it is rarely possible to record everything in detail, behavioral biologists use standardized sampling methods. Some common ones are:
Focal animal sampling
- One individual (the “focal animal”) is observed continuously for a set period.
- All behaviors of this individual, or all behaviors from a predefined subset, are recorded.
Use:
- Detailed study of an individual’s time budget (how it allocates time to activities).
- Tracking changes in one animal’s behavior across different situations.
Scan sampling
- At regular intervals (e.g., every minute), the observer quickly notes what each individual in a group is doing at that moment.
Use:
- Estimating the frequency of different behaviors in a group.
- Studying group-level patterns (e.g., proportion of animals feeding, resting, or being vigilant).
Behavior (or event) sampling
- Only specific behaviors of interest (e.g., aggressive acts, mating attempts) are recorded whenever they occur, regardless of which individual performs them.
Use:
- Studying relatively rare or brief events.
- Building up statistics on particular behavior types.
Recording Methods: When and How Behaviors Occur
Several standardized techniques help quantify behavior over time.
Continuous recording
- All occurrences of specified behaviors and their durations are recorded throughout the observation period.
Advantages:
- Provides detailed data on sequences and timing.
- Allows calculation of precise rates and durations.
Disadvantages:
- Very demanding for the observer.
- Often limited to a small set of behaviors.
Time sampling techniques
To make recording manageable, time is divided into intervals.
- Instantaneous (point) sampling:
- At each pre-set time point, the observer notes whether a behavior is occurring.
- Yields an estimate of the proportion of time spent in each behavior type.
- One-zero sampling:
- For each interval, the observer notes whether a behavior occurred at least once (1) or not (0).
- Suitable for frequent, brief behaviors that would be hard to record continuously.
These methods trade some precision for practicality but can still provide robust quantitative data when used carefully.
Use of Technology
Modern behavioral research often uses technical aids:
- Video and audio recordings for later detailed analysis and repeated viewing.
- Motion tracking (e.g., via GPS, RFID tags, or computer vision) to follow movements and interactions.
- Automated sensors (e.g., accelerometers, microphones) to monitor behavior in inaccessible environments or over long periods.
- Software tools for coding behaviors from recordings and performing statistical analyses.
These tools extend observational capabilities beyond what is possible with the naked eye and manual note-taking.
Experimental Methods
Experimental approaches involve controlled manipulations to test specific hypotheses about behavior.
Controlled Manipulation of Stimuli
Behavioral biologists often ask: “If X changes, does behavior Y change in a predictable way?” Common manipulations include:
- Stimulus intensity or quality:
- Varying loudness of calls, brightness of colors, or concentration of odors.
- Testing which aspects of a stimulus animals respond to.
- Stimulus presence/absence or arrangement:
- Adding or removing objects, conspecifics, or predators.
- Changing the configuration of cues in the environment (for navigation studies, for instance).
- Timing:
- Presenting stimuli at different times of day or life stages to examine sensitivity and internal states.
By comparing behavior across controlled conditions, researchers can infer causal relationships.
Control Groups and Experimental Design
To draw reliable conclusions, experiments must be designed carefully:
- Control groups receive no treatment (or a neutral treatment).
- Experimental groups are exposed to the treatment of interest.
Key considerations:
- Random assignment of individuals to groups where possible.
- Keeping other conditions constant (same light, temperature, feeding, etc.).
- Blinding observers so they do not know which animals are in which group, reducing bias.
Deprivation and Choice Experiments
Two common experimental types are:
Deprivation experiments
- Withholding specific experiences or stimuli to see if a behavior still appears.
- Used to explore the role of experience vs. innate predisposition in behavior.
Example frameworks (not detailed case studies):
- Raising animals without exposure to certain social cues to test whether specific courtship signals are learned or innate.
- Restricting access to particular landmarks to study learning in navigation.
Such experiments must take ethical considerations into account, particularly for social or cognitively complex animals.
Choice (preference) experiments
- Animals are given options (e.g., different foods, mates, habitats), and their choices are recorded.
- Used to analyze preferences and decision-making.
Design features:
- Present alternatives simultaneously in a controlled setup.
- Ensure that non-focal factors (e.g., lighting, position) do not bias choices.
Field Experiments and Semi-Natural Conditions
Not all experiments occur in the laboratory. Behavioral biologists also conduct:
- Field experiments directly in natural habitats, modifying aspects of the environment (e.g., adding model predators, altering resource distribution).
- Semi-natural experiments in large enclosures or controlled outdoor settings that mimic key aspects of natural conditions while allowing more control and observation.
These experiments bridge the gap between tightly controlled lab studies and fully natural observations.
Quantitative Analysis and Modeling
Behavior is variable and complex, so quantitative methods are essential.
Measuring and Summarizing Behavior
Common quantitative measures include:
- Frequency: how often a behavior occurs per unit time.
- Duration: how long behaviors last on average.
- Latency: time from stimulus onset to response.
- Intensity: graded measures of behavior strength (e.g., speed of movement, number of display elements).
Researchers summarize these measures with:
- Averages (means, medians).
- Variability (ranges, standard deviations).
- Proportions (e.g., proportion of time spent feeding vs. resting).
Statistical Testing
To determine whether observed differences are likely to reflect real effects rather than random variation, statistical tests are used:
- Comparing groups (e.g., treated vs. control).
- Testing correlations between variables (e.g., body size and dominance).
- Building models that incorporate multiple factors (e.g., age, sex, environmental conditions).
The choice of test depends on the type of data and study design, but the underlying goal is always the same: to assess whether patterns are statistically and biologically meaningful.
Modeling Behavior
Mathematical and computational models help:
- Predict how animals should behave under certain assumptions (e.g., maximizing energy gain).
- Simulate interactions within groups (e.g., collective movement, dominance hierarchies).
- Explore consequences of different strategies over evolutionary time.
These models are then tested against real behavioral data, creating a feedback loop between theory and observation.
Comparison, Interpretation, and Integration
Methods alone do not produce understanding; behavioral biologists must interpret results in broader contexts.
Comparative Studies
Comparisons can be made:
- Within species: between individuals, populations, or environments (e.g., behavior in urban vs. rural populations).
- Between species: especially among closely related species with different ecologies.
Goals:
- Identify general principles and exceptions.
- Distinguish between species-specific and widespread behaviors.
- Relate behavior to ecological and social conditions.
Integrating Multiple Levels of Explanation
Ultimately, behavioral biology aims to link:
- Mechanisms (physiology, neurobiology).
- Development (learning, ontogeny).
- Function (adaptive value).
- Evolution (phylogenetic history).
A comprehensive explanation of a behavior typically draws on all four levels. For example, a particular courtship display might be:
- Triggered by specific visual cues and hormones (mechanism).
- Refined through practice and social experience (development).
- Increasing mating success under given ecological conditions (function).
- Related to similar displays in sister species (evolution).
Careful use of the methods outlined above allows researchers to build such multi-layered explanations.
Ethical and Practical Considerations
Behavioral research involves living organisms and often sensitive social behaviors, so:
- Ethical guidelines govern the treatment of study animals.
- Minimizing stress and harm is a key requirement.
- Non-invasive and observational methods are preferred when possible.
- The potential benefits of research must justify any intrusions.
- In studies involving humans, informed consent, privacy protection, and special ethical oversight are essential.
Practical constraints (time, funding, access to study sites, animal availability) also shape which methods can be used and how extensive studies can be.
Summary
Behavioral biology seeks to:
- Describe behaviors systematically and objectively.
- Understand their immediate causes and mechanisms.
- Trace their development over an individual’s life.
- Evaluate their adaptive significance.
- Reconstruct their evolutionary history.
To achieve these goals, behavioral biologists employ:
- Structured observational methods (ethograms, sampling techniques).
- Controlled experiments (manipulating stimuli, using control groups).
- Quantitative analysis and modeling.
- Comparative and integrative approaches that connect behavior to physiology, development, ecology, and evolution.
These goals and methods form the foundation for the more specialized topics in behavioral biology that follow in later chapters.