DescriptionReading Guide: Lionfish Paper
This paper combines several topics we have been studying, including digestion and
metabolism, and previews our next unit, which relates to temperature’s effects on
physiology. In this paper, the author studies lionfish, which are invasive in the
Caribbean, to see how much energy the animal spends on digesting food (SDA) and how
water temperature affects both how much they eat and how fast they can digest it.
If you’re not used to reading scientific papers, don’t freak out. You’ll find that with some
patient attention and looking up a few words, you can actually understand most of this.
And more understanding will come when we work with the paper in class. Don’t start by
reading the Abstract (summary)—it’s too condensed to be easily understandable. Begin
with the Introduction.
As you encounter them in the paper, look up any words you don’t know. Note also the
list of abbreviations at the top of page 2! Let’s start with these:
1. Define each of the following, using your own words to do so as much as possible:
a) Standard Metabolic Rate (SMR) (and what is the difference between that and basal
metabolic rate, and between those and total metabolic rate?):
b) Postprandial:
c) SDA (not just what it stands for but what it means! See Ch 7 and Ch 7 goal sheet):
→ Review the ways that metabolic rate is measured in Chapter 7.
I. Introduction
The purpose of a scientific paper’s introduction is to explain why the study was worth
doing at all. Look for the major arguments the author is making.
2. The first thing they have to convince their readers is that it’s worth it to look at
metabolism in an invasive fish. What reason(s) do they give for doing this?
3. All animals have an SDA; that is, all animals spend energy to digest and absorb food
(see examples in Figure 1). So why is it “ecologically relevant” here—why bother looking
at it?
4. Next they explain why they are bothering to look at the effect of temperature. What
reason(s) do they give for looking at this? What do you personally imagine an effect of
higher temperature on feeding might be? Why?
They go on to discuss the role of metabolic phenotype (p.2) and feeding frequency (p.23), which are not our focus. Skip ahead to the first full paragraph on p. 3 where you can
read about lionfish themselves.
5. What is an interesting thing to you about lionfish, either from these two lionfish
paragraphs or an outside source, or (for those fish lovers out there?) your own
knowledge?
At the end of the Introduction, the authors give you their hypotheses, the ideas that
their experiments are going to test. Read these and note which ones do and don’t make
sense to you.
II. Materials and Methods
First they tell you how they caught the lionfish (“Animal Collection and Husbandry,”);
read or skim that.
Next they tell you how they measured metabolic rate.
6. As a reminder to yourself, why is measuring O2 consumption a good way to measure
how much energy an animal is using/burning (metabolic rate)?
The rest of the Methods can be skipped, except if you want to refer back later to see
exactly how they did something.
III. Results
7. How much more yummy fish and shellfish did lionfish eat in the environment that was
just 6 degrees C warmer (32 C vs. 26 C)? Also, how much did their metabolic rate (SMR)
go up between those two temps?
8. Carefully check out Figure 2. In your absolutely own words (avoid all the paper’s
words if possible; just do the best you can), what does it tell you?
9. Let’s look at parts of Figure 3. Start with panels I and M. We already saw in Figure 2
that the SDA peak (the highest amount of energy they spend metabolizing food) is
higher when the water temp is higher. What happens to the duration of the SDA (the
overall time it takes to process food) at warmer temps, and why do you think this
happens?
IV. Discussion
10. Based on the findings here as described in the first few paragraphs of the Discussion
section, it seems like lionfish have to lie on the couch and watch football after a big meal
at warm temps, because they have no remaining “scope for activity.” How can they get
away with using all their energy to digest food, without getting eaten themselves?
11. In the “Temperature” section of the Discussion, the authors explain why they think
the lionfish may benefit from a warming ocean. Why is that, and why might they benefit
more than other species of fish?
In the “Metabolic Phenotype” section, the authors note that lionfish seem to come in
different sorts or “personalities”– the active ones (high AMR, high scope for activity)
and the gluttons that are more of the focus here (more of their metabolic rate devoted
to SDA, leaving less available for activity). They suggest that this makes lionfish an even
more scary invasive species because one strategy will succeed in high-food
environments and the other in low-food environments, allowing the population as a
whole to survive in almost any environment. Scary! Or in other words, diversity within a
species is good!—but what’s good for an invasive species might be even worse for the
ecosystem.
12. This question exists so you read the paragraph above. Please go read the paragraph
above. Would you be willing to eat lionfish to help solve this invasive species problem?
© 2019. Published by The Company of Biologists Ltd | Journal of Experimental Biology (2019) 222, jeb209437. doi:10.1242/jeb.209437
RESEARCH ARTICLE
An appetite for invasion: digestive physiology, thermal
performance and food intake in lionfish (Pterois spp.)
ABSTRACT
Species invasions threaten global biodiversity, and physiological
characteristics may determine their impact. Specific dynamic action
(SDA; the increase in metabolic rate associated with feeding and
digestion) is one such characteristic, strongly influencing an animal’s
energy budget and feeding ecology. We investigated the relationship
between SDA, scope for activity, metabolic phenotype, temperature
and feeding frequency in lionfish (Pterois spp.), which are invasive to
western Atlantic marine ecosystems. Intermittent-flow respirometry
was used to determine SDA, scope for activity and metabolic
phenotype at 26°C and 32°C. Maximum metabolic rate occurred
during digestion, as opposed to exhaustive exercise, as in more
athletic species. SDA and its duration (SDAdur) were 30% and 45%
lower at 32°C than at 26°C, respectively, and lionfish ate 42% more at
32°C. Despite a 32% decline in scope for activity from 26°C to 32°C,
aerobic scope may have increased by 24%, as there was a higher
range between standard metabolic rate (SMR) and peak SDA
(SDApeak; the maximum postprandial metabolic rate). Individuals
with high SMR and low scope for activity phenotypes had a less costly
SDA and shorter SDAdur but a higher SDApeak. Feeding frequently
had a lower and more consistent cost than consuming a single meal,
but increased SDApeak. These findings demonstrate that: (1) lionfish
are robust physiological performers in terms of SDA and possibly
aerobic scope at temperatures approaching their thermal maximum,
(2) lionfish may consume more prey as oceans warm with climate
change, and (3) metabolic phenotype and feeding frequency may be
important mediators of feeding ecology in fish.
KEY WORDS: Invasive species, Metabolic phenotype,
Specific dynamic action, Thermal physiology, Temperature
INTRODUCTION
Species invasions are a widely recognized and growing threat to
global biodiversity (Ricciardi et al., 2017). Dozens of fish species
have become invasive worldwide as the result of human activity,
and many more may become invasive, with over 600 known
introductions of non-native species (Gozlan, 2008). The probability
of an introduced species becoming invasive depends on ecological
and life history traits (e.g. niche, growth rate, reproductive capacity
1
Fish Ecology and Conservation Physiology Lab, Carleton University, 1125 Colonel
By Drive, Ottawa, ON, Canada, K1S 5B6. 2The Cape Eleuthera Institute, Eleuthera,
The Bahamas. 3Fisheries and Oceans Canada, 80 East White Hills Road,
PO Box 5667, St John’s, NL, Canada, A1C 5X1. 4Department of Ecology, Evolution
and Marine Biology, University of California, Santa Barbara, Santa Barbara,
CA 93106, USA.
*Author for correspondence (clay.steell@carleton.ca)
S.C.S., 0000-0001-7003-145X; T.E.V.L., 0000-0003-0209-543X; J.W.B., 00000003-2496-8764; S.J.C., 0000-0002-5407-0659; E.J.E., 0000-0002-0120-7498
Received 23 June 2019; Accepted 9 September 2019
or tolerance to disturbance), which are underpinned by
physiological characteristics (Van Kleunen et al., 2010; Kelley,
2014; Lennox et al., 2015). Characteristics of metabolic rate – the
rate at which an organism expends energy, commonly measured
using oxygen consumption rate (M_ O2 ) – are increasingly recognized
as a potential determinant of invasion success (González-Ortegón
et al., 2010; Maazouzi et al., 2011; Lejeusne et al., 2014; Lagos
et al., 2017). Relatively few studies have examined their relevance to
invasive fish, however, which is surprising given the number and
impact of invasive fish species worldwide (but see Marras et al.,
2015; McCallum et al., 2017; Behrens et al., 2017; Srean et al.,
2017; Nati et al., 2018; Tessier et al., 2018).
A characteristic of metabolic rate that has largely been ignored in
invasive species biology is specific dynamic action (SDA): the
postprandial increase in metabolic rate associated with feeding and
digestion. SDA represents the total energy expenditure of numerous
pre-absorptive, absorptive and post-absorptive pathways associated
with feeding and digestion, and is typified by a rapid rise in
metabolic rate up to a relatively short-lived peak (SDApeak) followed
by a longer decline (Jobling, 1981; McCue, 2006; Chabot et al.,
2016) (Fig. 1). SDA is ecologically relevant because it can occupy a
large proportion of a fish’s energy budget in terms of its aerobic
scope or scope for activity. Aerobic scope is the range between
standard metabolic rate (SMR, the minimum metabolic rate required
for maintenance) and maximum metabolic rate (MMR), whereas
scope for activity is the range between SMR and active metabolic
rate (AMR, the metabolic rate elicited from maximal exercise)
(Sandblom et al., 2014; Norin and Clark, 2016). Aerobic scope and
scope for activity are equivalent and interchangeable in many
species (i.e. AMR=MMR); however, in some more-sedentary
species, these characteristics are distinct as MMR may be achieved
independently of exhaustive exercise (i.e. AMR
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