Beyond Memorization: Exercises that Help Students Forge, Remember, and Apply their Knowledge.

This personal retrospective describes what I consider to be the two most important learning practices in my classroom: A form of interactive engagement that I call collaborative exercises and general principles exercises. The power of interactive engagement had already been described and evaluated in the pedagogical literature of the physical sciences, but was not yet used widely in biology when I began teaching. Here, I discuss practical suggestions for implementing IE successfully and strategies for inviting students to try a new way of learning. General principles are everywhere we look-in lectures, titles of textbooks, and in our own minds as we process new information. Many of our students, however, are only marginally familiar with how general principles can revolutionize the way they learn, remember information, and solve novel context problems. The general principles exercise stimulates students not only to engage actively with received general principles from lecture, reading, and other sources, but also to develop the transferable skill of discovering (not just being told about) the patterns that permeate the natural world and our study of it.

Torpor and hypothermia: reversed hysteresis of metabolic rate and body temperature.

Regulated torpor and unregulated hypothermia are both characterized by substantially reduced body temperature (Tb) and metabolic rate (MR), but they differ physiologically. Although the remarkable, medically interesting adaptations accompanying torpor (e.g., tolerance for cold and ischemia, absence of reperfusion injury, and disuse atrophy) often do not apply to hypothermia in homeothermic species such as humans, the terms "torpor" and "hypothermia" are often used interchangeably in the literature. To determine how these states differ functionally and to provide a reliable diagnostic tool for differentiating between these two physiologically distinct states, we examined the interrelations between Tb and MR in a mammal (Sminthopsis macroura) undergoing a bout of torpor with those of the hypothermic response of a similar-sized juvenile rat (Rattus norvegicus). Our data show that under similar thermal conditions, 1) cooling rates differ substantially (approximately fivefold) between the two states; 2) minimum MR is approximately sevenfold higher during hypothermia than during torpor despite a similar Tb; 3) rapid, endogenously fuelled rewarming occurs in torpor but not hypothermia; and 4) the hysteresis between Tb and MR during warming and cooling proceeds in opposite directions in torpor and hypothermia. We thus demonstrate clear diagnostic physiological differences between these two states that can be used experimentally to confirm whether torpor or hypothermia has occurred. Furthermore, the data can clarify the results of studies investigating the ability of physiological or pharmacological agents to induce torpor. Consequently, we recommend using the terms "torpor" and "hypothermia" in ways that are consistent with the underlying regulatory differences between these two physiological states.

Dietary lipid saturation influences environmental temperature preference but not resting metabolic rate in the Djungarian Hamster (Phodopus sungorus).

Heterothermic rodents increase self-selection of diets rich in polyunsaturated fatty acids (PUFAs) when exposed to cold, short days, or short-day melatonin profiles, and Djungarian hamsters (Phodopus sungorus) do so in long days in response to cold exposure alone. To determine whether Djungarian hamsters are also capable of selecting a thermal environment in response to dietary lipid composition, continuously normothermic hamsters were fed either a PUFA-rich diet or a diet rich in saturated fatty acids (SFAs) for 6-10 wk and given a choice of thermal environments. As predicted, SFA-fed hamsters were more likely than PUFA-fed hamsters to occupy the single heated corner of their cage ([Formula: see text]) and were most likely to show this diet-related difference in behavior when T(a) fell within the thermal neutral zone. Respirometry revealed no effect of diet on whole-animal or mass-specific resting metabolic rate or on lower critical temperature. The results are more consistent with the homeoviscous adaptation hypothesis, which predicts that organisms should make physiological and/or behavioral adjustments that preserve membrane fluidity within a relatively small range, than with the membrane pacemaker hypothesis, which predicts that high PUFA content in membrane phospholipids should increase basal metabolic rate.

Gut bacteria profiles of Mus musculus at the phylum and family levels are influenced by saturation of dietary fatty acids.

BACKGROUND: Mammalian gut microbiota have been implicated in a variety of functions including the breakdown of ingested nutrients, the regulation of energy intake and storage, the control of immune system development and activity, and the synthesis of novel chemicals. Previous studies have shown that feeding mammalian hosts a high-fat diet shifts gut bacteria at the phylum level to reduce the ratio of Bacteroidetes-to-Firmicutes, while feeding hosts a fat-restricted diet increases this ratio. However, few studies have investigated the differential effects of fatty acid type on gut bacterial profile. METHODS: Over a 14-week period, Mus musculus were fed a diet rich in omega-3 polyunsaturated fatty acids (n-3 PUFAs), omega-6 polyunsaturated fatty acids (n-6 PUFAs), or saturated fatty acids (SFAs). Fecal pellets were collected before and after the treatment period from 12 randomly selected mice (4 per treatment group). Bacterial DNA was extracted from the pellets and characterized by analysis of the hypervariable V3 region of the 16S rRNA. Nominal logistic regression models were used to assess shifts in microbial profile at the phylum and family levels in response to diet. RESULTS: A significant decrease in the proportion of phylum Bacteroidetes species was observed for mice fed any of the three diets over time. However, the SFA-rich diet group showed a significantly greater decrease in Bacteroidetes proportion (-28%) than did either the n-3 PUFA group (-10%) or the n-6 PUFA group (-12%). At the family level, a significant decrease in proportion of Porphyromonadaceae was observed for mice fed the n-6 PUFA-rich diet, and a significant decrease in proportion of Lachnospiraceae was observed for mice fed the SFA-rich diet. There was no significant effect of diet type on body mass change. CONCLUSION: Our results indicate that SFAs have stronger effects than PUFAs in shifting gut microbiota profiles toward those typical of obese individuals, and that dietary fatty acid saturation influences shifts in gut microbiota independently of changes in body mass.

Teaching simple experimental design to undergraduates: do your students understand the basics?

This article provides instructors with guidelines for teaching simple experimental design for the comparison of two treatment groups. Two designs with specific examples are discussed along with common misconceptions that undergraduate students typically bring to the experiment design process. Features of experiment design that maximize power and minimize the effects of interindividual variation, thus allowing reduction of sample sizes, are described. Classroom implementation that emphasizes student-centered learning is suggested, and thought questions, designed to help students discover and name the basic principles of simple experiment design for themselves, are included with an answer key.

The strong-inference protocol: not just for grant proposals.

The strong-inference protocol puts into action the important concepts in Platt's often-assigned, classic paper on the strong-inference method (10). Yet, perhaps because students are frequently performing experiments with known outcomes, the protocols they write as undergraduates are usually little more than step-by-step instructions for performing the experiment. The strong-inference protocol, however, includes an explicit statement of possible experimental outcomes and the interpretation that would follow from each. This approach encourages thorough planning, enhances the efficiency of experimental designs, and increases the power of statistical analysis by explicitly stating a priori predictions as well as the statistical methods that will be used to test them. A sample protocol for an experiment investigating temperature-metabolism relations in chicken embryos is provided to illustrate the important components of the strong-inference protocol and to encourage instructors to incorporate this powerful research tool into undergraduate laboratory courses.

Are chicken embryos endotherms or ectotherms? A laboratory exercise integrating concepts in thermoregulation and metabolism.

This investigative laboratory exercise uses the different relations between ambient temperature and metabolic rate in endotherms and ectotherms as a core concept to answer the following question: What thermoregulatory mode is employed by chicken embryos? Emphasis is placed on the physiological concepts that can be taught with this exercise, including methods for measuring rates of oxygen consumption, the relation between oxygen consumption and metabolic rate, the influence of temperature on metabolic rate, and the differences between endotherms and ectotherms both in the overall magnitude of metabolic rate and in the shape of the relation between metabolic rate and ambient temperature. Included in this article are respirometer designs suitable for teachers working with a wide variety of budgets and available equipment, specific laboratory protocols for collecting data, sample data, thought questions with sample answers, and suggestions for classroom implementation as a 1-, 2- or 3-wk laboratory exercise that can be taught at a variety of undergraduate levels.

Photoperiod affects daily torpor and tissue fatty acid composition in deer mice.

Photoperiod and dietary lipids both influence thermal physiology and the pattern of torpor of heterothermic mammals. The aim of the present study was to test the hypothesis that photoperiod-induced physiological changes are linked to differences in tissue fatty acid composition of deer mice, Peromyscus maniculatus ( approximately 18-g body mass). Deer mice were acclimated for >8 weeks to one of three photoperiods (LD, light/dark): LD 8:16 (short photoperiod), LD 12:12 (equinox photoperiod), and LD 16:8 (long photoperiod). Deer mice under short and equinox photoperiods showed a greater occurrence of torpor than those under long photoperiods (71, 70, and 14%, respectively). The duration of torpor bouts was longest in deer mice under short photoperiod (9.3 +/- 2.6 h), intermediate under equinox photoperiod (5.1 +/- 0.3 h), and shortest under long photoperiod (3.7 +/- 0.6 h). Physiological differences in torpor use were associated with significant alterations of fatty acid composition in approximately 50% of the major fatty acids from leg muscle total lipids, whereas white adipose tissue fatty acid composition showed fewer changes. Our results provide the first evidence that physiological changes due to photoperiod exposure do result in changes in lipid composition in the muscle tissue of deer mice and suggest that these may play a role in survival of low body temperature and metabolic rate during torpor, thus, enhancing favourable energy balance over the course of the winter.

Daily timed melatonin feedings mimic effects of short days on testis regression and cortisol in circulation in Siberian hamsters.

This study tested the efficacy of timed oral administration of melatonin as an alternative both to invasive methods (daily injections, timed infusions) and to untimed oral administration in Siberian hamsters (Phodopus sungorus), an important model for the study of photoperiodism. Hamsters readily consumed a small piece of melatonin-treated apple immediately when presented and circulating melatonin was rapidly elevated with a half-life of approximately 3.5 h. Melatonin-treated apple was fed to hamsters for 3 weeks at 2 h before lights off to extend the duration of the nighttime rise in endogenous melatonin. Melatonin treatment induced testicular regression and elevated serum cortisol, effects comparable to those in hamsters exposed to short days. These findings support the hypothesis that timed oral administration of melatonin can mimic the effects of short days and provide a method by which melatonin can be delivered without the potentially confounding effects of handling and injection stress.

Fitness correlates of song repertoire size in free-living song sparrows (Melospiza melodia).

Models of sexual selection propose that exaggerated secondary sexual ornaments indicate a male's own fitness and the fitness of his offspring. These hypotheses have rarely been thoroughly tested in free-living individuals because overall fitness, as opposed to fitness components, is difficult to measure. We used 20 years of data from song sparrows (Melospiza melodia) inhabiting Mandarte Island, British Columbia, Canada, to test whether a male's song repertoire size, a secondary sexual trait, predicted overall measures of male or offspring fitness. Males with larger song repertoires contributed more independent and recruited offspring, and independent and recruited grandoffspring, to Mandarte's population. This was because these males lived longer and reared a greater proportion of hatched chicks to independence from parental care, not because females mated to males with larger repertoires laid or hatched more eggs. Furthermore, independent offspring of males with larger repertoires were more likely to recruit and then to leave more grandoffspring than were offspring of males with small repertoires. Although we cannot distinguish whether observed fitness variation reflected genetic or environmental effects on males or their offspring, these data suggest that female song sparrows would gain immediate and intergenerational fitness benefits by pairing with males with large song repertoires.

Simulated human diving and heart rate: making the most of the diving response as a laboratory exercise.

Laboratory exercises in which students examine the human diving response are widely used in high school and college biology courses despite the experience of some instructors that the response is unreliably produced in the classroom. Our experience with this exercise demonstrates that the bradycardia associated with the diving response is a robust effect that can easily be measured by students without any sophisticated measurement technology. We discuss measures that maximize the success of the exercise by reducing individual variation, designing experiments that are minimally affected by change in the response over time, collecting data in appropriate time increments, and applying the most powerful statistical analysis. Emphasis is placed on pedagogical opportunities for using this exercise to teach general principles of physiology, experimental design, and data analysis. Data collected by students, background information for instructors, a discussion of the relevance of the diving reflex to humans, suggestions for additional experiments, and thought questions with sample answers are included.

Djungarian hamsters exhibit temperature-dependent dietary fat choice in long days.

Previous studies have shown that whole-animal thermal responses of ectotherms and heterotherms (e.g., hibernators), both of which experience a wide range of body temperatures, are related to the saturation level of somatic lipids, which in turn can be influenced by the ratio of saturated and unsaturated fatty acids in the diet. This study demonstrates that Djungarian hamsters held in long days display ambient temperature-dependent choice of dietary fats, increasing their preference for saturated fats when ambient temperature increases (to 27 degrees C) and later reversing this preference when ambient temperature is returned to its original value (8 degrees C). Changes in percent contribution of the unsaturated and saturated diets in response to temperature were accomplished almost solely by changes in the amount of unsaturated diet consumed. Temperature-dependent fatty acid choice occurs at a stage in the annual cycle when Djungarian hamsters do not enter spontaneous daily torpor and therefore experience only small changes in core body temperature. These results suggest that temperature-dependent fatty acid choice may occur in a wide range of animals, including nonheterothermic endotherms.

Site-fidelity, longevity, and population dynamics of broad-tailed hummingbirds: a ten year study.

In the course of other studies we have amassed a decade of records from banding, and observing the nests of, a breeding population of broad-tailed hummingbirds in Colorado, USA. In addition we have less extensive banding records for two other hummingbird species that migrate through the area but do not breed there. The rate of return of broad-tails between consecutive breeding seasons has been as high as 70% for females and 27% for males, suggesting substantial site fidelity and malebiased dispersal or mortality; our records also suggest that rufous hummingbirds are faithful to a particular migratory route. The oldest recaptured birds were at least eight years old, an age that exceeds predictions based on allometric extrapolation from other bird species; the apparent yearly survival rate of females is also unexpectedly high for birds of such small body mass. The earliest broad-tails to arrive at the start of the breeding season appear to be older, experienced individuals. Reuse of a specific nest site between consecutive years by the same female or different females seems to depend on the success of nesting efforts at that site. We have calculated the rate of change in size of the broad-tail population based on our estimates of female survivorship and fledging success. By this method the population appears to be declining, although nest counts themselves suggest that numbers of breeding females have remained fairly constant at least over the last seven years of our study.