The Jellyfish Cassiopea Exhibits a Sleep-like State
Open accessAbstract
Do all animals sleep? Sleep has been observed in many vertebrates, and there is a growing body of evidence for sleep-like states in arthropods and nematodes [1-5]. Here we show that sleep is also present in Cnidaria [6-8], an earlier-branching metazoan lineage. Cnidaria and Ctenophora are the first metazoan phyla to evolve tissue-level organization and differentiated cell types, such as neurons and muscle [9-15]. In Cnidaria, neurons are organized into a non-centralized radially symmetric nerve net [11, 13, 15-17] that nevertheless shares fundamental properties with the vertebrate nervous system: action potentials, synaptic transmission, neuropeptides, and neurotransmitters [15-20]. It was reported that cnidarian soft corals [21] and box jellyfish [22, 23] exhibit periods of quiescence, a pre-requisite for sleep-like states, prompting us to ask whether sleep is present in Cnidaria. Within Cnidaria, the upside-down jellyfish Cassiopea spp. displays a quantifiable pulsing behavior, allowing us to perform long-term behavioral tracking. Monitoring of Cassiopea pulsing activity for consecutive days and nights revealed behavioral quiescence at night that is rapidly reversible, as well as a delayed response to stimulation in the quiescent state. When deprived of nighttime quiescence, Cassiopea exhibited decreased activity and reduced responsiveness to a sensory stimulus during the subsequent day, consistent with homeostatic regulation of the quiescent state. Together, these results indicate that Cassiopea has a sleep-like state, supporting the hypothesis that sleep arose early in the metazoan lineage, prior to the emergence of a centralized nervous system.
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TL;DR
The upside-down jellyfish Cassiopea shows a nighttime quiescent state with reduced responsiveness and rebound after deprivation, meeting behavioral criteria for a sleep-like state. This suggests sleep-like behavior may predate centralized nervous systems.
Key findings
- Across 23 jellyfish tracked over six day-night cycles, nighttime pulsing dropped about 32% relative to daytime: 781 ± 199 vs 1155 ± 315 pulses per 20 min. A separate 16-jellyfish no-feeding cohort showed the same pattern.
- During night, latency to first pulse after a drop was 5.9 ± 4.0 s vs 2.1 ± 0.9 s during day; time to reach bottom was 12.0 ± 3.2 s vs 8.6 ± 2.9 s during day (n = 23). A second drop within 30 s abolished the day-night difference, indicating rapid reversibility.
- Depriving jellyfish of nighttime quiescence for the last 6 h reduced next-day first-4-h activity by ~12% (1146 ± 232 to 1008 ± 210 pulses/20 min, n = 30); 12-h deprivation reduced following-day activity by ~17% (1361 ± 254 to 1132 ± 263, n = 16). The same mechanical stimulation during the day produced no significant change.
- Constant light did not produce circadian oscillation in activity, but constant dark did; melatonin and pyrilamine induced concentration-dependent daytime quiescence.
Methods
- Behavioral tracking: individual Cassiopea in clear cubbies inside light-controlled tanks under 12:12 light/dark; overhead cameras at 15 fps recorded pulsing; pulse events and inter-pulse intervals were derived from pixel-intensity changes in manually selected regions of interest.
- Samples/design: 23 jellyfish tracked for 6 consecutive day-night cycles; 16 tracked for 3 cycles without feeding; arousal assay n = 23; 6-h and 12-h deprivation experiments n = 30 and n = 16; additional clonal C. xamachana used for some pharmacology/circadian assays. Genotyping via COI sequencing and actin staining/confocal imaging were also performed.
- Analysis: SciPy-based processing; paired/unpaired t-tests, two-way ANOVA with Bonferroni posttest; normality and equal-variance checks; log transformation for time-to-first-pulse data.
Limitations
- Stated: the authors note that whether other cnidarians sleep remains untested, and the ancestral function/selective advantage of sleep is unresolved. They also state no sample-size predetermination, no blinding during experiments/outcome assessment, and no specific randomization method.
- Evident: most experiments used a mixed collection of Cassiopea species/sizes/ages/sexes/morphologies, so species-level generalization is limited; some assays used a young clonal C. xamachana line.
- Evident: the sleep-like state is inferred from behavioral proxies (pulsing, latency, rebound) under laboratory conditions; circadian regulation is ambiguous because activity cycled in constant dark but not under several constant-light conditions, and mechanistic evidence is limited to pharmacological responses.
Model-written summary. Check it against the paper before relying on it.
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Claims the paper makes
0 of 39 cited sources checked · 0 supported · 0 flagged
- Findingown data
Melatonin induces a reversible, concentration-dependent decrease in Cassiopea activity during the day.
“We observed that melatonin induces a reversible decrease in activity in Cassiopea during the day in a concentration-dependent manner (Figure S4F–H), suggesting that melatonin has a conserved quiescence-inducing effect in Cassiopea.”
Quote in the paperSubstantial claim- Rests on an assumption: Pulsing frequency is a valid measure of behavioral activity and quiescence in Cassiopea.
- Findingown data
A 6-hour mechanical perturbation during the last 6 hours of night caused a significant ~12% decrease in Cassiopea activity during the first 4 hours of the following day compared with the pre-perturbation day.
“When the perturbation was performed during the last 6 hours of the night (Figure 4A), we observed a significant decrease in activity (~12%) during the first 4 hours of the following day relative to the pre-perturbation day (mean of first 4 hours of pre-perturbation day: 1146 ± 232 pulses/20 min compared to post-perturbation day: 1008 ± 210 pulses/20 min, mean ± s.d.; n = 30 animals; Figure 4C).”
Quote in the paperSubstantial claim- Rests on an assumption: Pulsing frequency is a valid measure of behavioral activity and quiescence in Cassiopea.
- Findingown data
Pyrilamine induces concentration-dependent quiescence in Cassiopea.
“Pyrilamine, a histamine H1 receptor antagonist that induces sleep in vertebrates [36], also induces concentration-dependent quiescence in Cassiopea (Figure S4F).”
Quote in the paperSubstantial claim- Rests on an assumption: Pulsing frequency is a valid measure of behavioral activity and quiescence in Cassiopea.
- Findingown data
During the decreased-activity periods after either 6-hour or 12-hour nighttime perturbation, Cassiopea showed increased response latency to a sensory stimulus.
“During periods of decreased activity after either the 6-hour or 12-hour perturbation, we also observed increased response latency to a sensory stimulus (Figure S4A), indicating a sleep-like state.”
Quote in the paperSubstantial claim- Rests on an assumption: Pulsing frequency is a valid measure of behavioral activity and quiescence in Cassiopea.
- Findingown data
No circadian oscillation of Cassiopea activity was observed under constant light conditions of low, mid, or full intensity.
“We observed no circadian oscillation of jellyfish activity under any of the constant light conditions (Figure S4D).”
Quote in the paperSubstantial claim- Rests on an assumption: Pulsing frequency is a valid measure of behavioral activity and quiescence in Cassiopea.
- Findingown data
Circadian oscillation of Cassiopea activity was observed under constant dark conditions.
“However, we do observe circadian oscillation of activity in constant dark conditions (Figure S4E).”
Quote in the paperSubstantial claim- Rests on an assumption: Pulsing frequency is a valid measure of behavioral activity and quiescence in Cassiopea.
- Findingown data
A brief 10 s water pulse every 20 min causes a transient increase in Cassiopea pulsing activity lasting about 5 min.
“The stimulus consisted of a brief (10 s) pulse of water every 20 min, which caused a transient increase in pulsing activity (Movie S2). This increase in pulsing activity lasts for approximately 5 min after the 10 s pulse of water.”
Quote in the paperSubstantial claim- Rests on an assumption: Pulsing frequency is a valid measure of behavioral activity and quiescence in Cassiopea.
- Findingown data
After a single day of decreased activity, Cassiopea return to baseline day and night activity levels.
“After a single day of decreased activity, the jellyfish return to baseline levels of day and night activity.”
Quote in the paperSubstantial claim- Rests on an assumption: Pulsing frequency is a valid measure of behavioral activity and quiescence in Cassiopea.
- Findingown data
Individual Cassiopea jellyfish showed different basal activity levels.
“While individual jellyfish showed different basal activity levels (Figure 2C)”
Quote in the paperNot a substantial claim- Rests on an assumption: Pulsing frequency is a valid measure of behavioral activity and quiescence in Cassiopea.
- Findingreasoning
When released into the water column, Cassiopea quickly reorient and move to the bottom of the tank.
“If Cassiopea are released into the water column, they quickly reorient and move to the bottom of the tank.”
Quote in the paperSubstantial claim - Findingown data
Despite variation in basal activity, the relative day-to-night change in activity was similar among jellyfish.
“Despite variations in basal activity, the relative change from day to night was similar between jellyfish (Figure 2D).”
Quote in the paperNot a substantial claim- Rests on an assumption: Pulsing frequency is a valid measure of behavioral activity and quiescence in Cassiopea.
- Findingown data
Cassiopea pulsing activity peaked upon feeding, on the fourth hour of each day.
“Pulsing activity peaked upon feeding, occurring on the 4th hour of each day (Figure 2C,D).”
Quote in the paperNot a substantial claim- Rests on an assumption: Pulsing frequency is a valid measure of behavioral activity and quiescence in Cassiopea.
- Premiseown data
Cassiopea continuously pulse by relaxing and contracting their bell at about 1 pulse per second.
“Cassiopea continuously pulse by relaxing and contracting their bell at a rate of about 1 pulse/second (Figure 1D).”
Quote in the paperSubstantial claim - Interpretationreasoning
Sleep arose early in the metazoan lineage, before the emergence of a centralized nervous system.
“Together these results indicate that Cassiopea has a sleep-like state, supporting the hypothesis that sleep arose early in the metazoan lineage, prior to the emergence of a centralized nervous system.”
- Also stated in: § RESULTS AND DISCUSSION · hedged
“suggesting that this behavioral state arose prior to the evolution of a centralized nervous system.”
Quote in the paperSubstantial claim- Rests on an assumption: Laboratory-recorded pulsing behavior reflects the jellyfish's natural behavioral state.
- Also stated in: § RESULTS AND DISCUSSION · hedged
- Interpretationreasoning
The ancestral role of sleep is rooted in basic requirements conserved across the animal kingdom.
“Our discovery of a sleep-like state in an ancient metazoan phylum suggests that the ancestral role of sleep is rooted in basic requirements that are conserved across the animal kingdom.”
Quote in the paperNot a substantial claim- Rests on an assumption: Laboratory-recorded pulsing behavior reflects the jellyfish's natural behavioral state.
- Interpretationown data
At least some mechanisms involved in vertebrate sleep may be conserved in Cassiopea.
“These results suggest that at least some mechanisms involved in vertebrate sleep may be conserved in Cassiopea.”
Quote in the paperSubstantial claim - Interpretationcitation
The observation of behaviorally and mechanistically conserved sleep-like states across animals strongly supports an early rooted sleep state rather than many instances of convergent evolution.
“The observation of behaviorally and mechanistically conserved sleep-like states across the animal kingdom [6,7] strongly supports the possibility for an early rooted sleep state rather than many instances of convergent evolution.”
Quote in the paperSubstantial claim0 of 2 sources checked - Interpretationown data
The quiescent state of Cassiopea may be under circadian regulation.
“This result suggests that the quiescent state may be under circadian regulation.”
Quote in the paperSubstantial claim - Interpretationcitation
A sleep-like state may not be specific to Cassiopea; other cnidarians may also exhibit it.
“Although future studies are required to test whether other cnidarians sleep, field studies showing behavioral quiescence, diel vertical migration, and swimming speeds that vary with diel period [21,22,23,37] suggest that a sleep-like state may not be specific to Cassiopea.”
Quote in the paperNot a substantial claim0 of 4 sources checked- cites [21] Kremien, 2013, Benefit of pulsation in soft corals, Proc. Natl. Acad. Sci. USA source
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- cites [22] Garm, 2012, Opposite patterns of diurnal activity in the box jellyfish Tripedalia cystophora and Copula sivickisi, Biol. Bull. source
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- cites [23] Seymour, 2004, Do box jellyfish sleep at night?, Med. J. Aust. source
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- cites [37] Moriarty, 2012, Vertical and horizontal movement patterns of scyphozoan jellyfish in a fjord-like estuary, Mar. Ecol. Prog. Ser. source
Not checked in the free run.
- Rests on an assumption: Laboratory-recorded pulsing behavior reflects the jellyfish's natural behavioral state.
- Interpretationown data
Cassiopea has a sleep-like state.
“Together these results indicate that Cassiopea has a sleep-like state, supporting the hypothesis that sleep arose early in the metazoan lineage, prior to the emergence of a centralized nervous system.”
- Also stated in: § RESULTS AND DISCUSSION · plain
“Cassiopea display the key behavioral characteristics of a sleep-like state: a reversible quiescent state with reduced responsiveness to stimuli and both homeostatic and possibly circadian regulation.”
Quote in the paperSubstantial claim- Rests on an assumption: Pulsing frequency is a valid measure of behavioral activity and quiescence in Cassiopea.
- Rests on an assumption: Laboratory-recorded pulsing behavior reflects the jellyfish's natural behavioral state.
- Also stated in: § RESULTS AND DISCUSSION · plain
- Interpretationown data
Cassiopea exhibit a nighttime quiescent state.
“These results demonstrate that Cassiopea have a quiescent state during the night.”
- Also stated in: § RESULTS AND DISCUSSION · hedged
“Cassiopea display the key behavioral characteristics of a sleep-like state: a reversible quiescent state with reduced responsiveness to stimuli and both homeostatic and possibly circadian regulation.”
Quote in the paperSubstantial claim- Rests on an assumption: Pulsing frequency is a valid measure of behavioral activity and quiescence in Cassiopea.
- Also stated in: § RESULTS AND DISCUSSION · hedged
- Interpretationreasoning
The stationary pulsing behavior makes Cassiopea suitable for behavioral tracking.
“This stationary pulsing behavior makes Cassiopea a suitable jellyfish for behavioral tracking.”
Quote in the paperSubstantial claim - Interpretationown data
Cassiopea have a nighttime quiescent state that is homeostatically controlled.
“Taken together, these results demonstrate that Cassiopea have a nighttime-quiescent state that is homeostatically controlled.”
Quote in the paperSubstantial claim- Rests on an assumption: Pulsing frequency is a valid measure of behavioral activity and quiescence in Cassiopea.
- Interpretationcitation
The finding is the first example of a sleep-like state in an organism with a diffuse nerve net.
“To our knowledge, our finding is the first example of a sleep-like state in an organism with a diffuse nerve net [7,8], suggesting that this behavioral state arose prior to the evolution of a centralized nervous system.”
Quote in the paperSubstantial claim0 of 2 sources checked - Interpretationreasoning
The mechanical perturbation disrupts quiescence for about 25% of the perturbation period.
“Thus, the perturbation disrupts quiescence for approximately 25% of the perturbation period (either 6 hours or 12 hours).”
Quote in the paperNot a substantial claim - Premisecitation
Many aspects of the nervous system are conserved between cnidarians and bilaterians, including neuropeptides and neurotransmitters.
“Though at least 600 million years of evolution separate cnidarians from bilaterians [9–16], many aspects of the nervous system are conserved, including neuropeptides and neurotransmitters [15–20].”
Quote in the paperSubstantial claim0 of 6 sources checked- cites [15] Bosch, 2017, Back to the basics: cnidarians start to fire, Trends Neurosci. source
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- cites [16] Grimmelikhuijzen, 1995, The nervous systems of cnidarians, EXS
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- cites [17] Satterlie, 2011, Do jellyfish have central nervous systems?, J. Exp. Biol. source
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- cites [18] Watanabe, 2009, Cnidarians and the evolutionary origin of the nervous system, Dev. Growth Differ. source
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- cites [19] Dupre, 2017, Non-overlapping neural networks in Hydra vulgaris, Curr. Biol. source
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- cites [20] Grimmelikhuijzen, 2002, Neuropeptides in cnidarians, Can. J. Zool. source
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- May combine two claims. 18
- Classified as probably not a substantial claim. 15
May combine two claims.
Findingown dataCassiopea show reduced responsiveness to a sensory stimulus at night, with longer time to first pulse and time to reach bottom compared with day.
“At night, the jellyfish showed an increase in the time to first pulse and the time to reach bottom, compared to day (time to first pulse day: 2.1 ± 0.9 s versus night: 5.9 ± 4.0 s, and the time to reach bottom day: 8.6 ± 2.9 s versus night: 12.0 ± 3.2 s, mean ± s.d.; n = 23 animals) (Figure 3B,C).”
- Also stated in: § RESULTS AND DISCUSSION · hedged
“This increased latency in response to stimulus indicates that Cassiopea have reduced responsiveness to stimulus during the night.”
Quote in the paperSubstantial claim- Rests on an assumption: Pulsing frequency is a valid measure of behavioral activity and quiescence in Cassiopea.
- Also stated in: § RESULTS AND DISCUSSION · hedged
May combine two claims.
Findingown dataThe nighttime quiescent state of Cassiopea is rapidly reversible, as a nighttime food stimulus transiently raised activity to daytime levels.
“To test the reversibility of this nighttime quiescent state we introduced a food stimulus at night, which transiently increased activity to daytime levels (Figure S3E).”
- Also stated in: § RESULTS AND DISCUSSION · plain
“The nighttime quiescent state in Cassiopea is thus rapidly reversible, consistent with a sleep-like behavior.”
Quote in the paperSubstantial claim- Rests on an assumption: Pulsing frequency is a valid measure of behavioral activity and quiescence in Cassiopea.
- Also stated in: § RESULTS AND DISCUSSION · plain
May combine two claims.
Findingown dataDaytime mechanical perturbation for 6 or 12 hours produced no significant pre- versus post-perturbation activity difference, indicating that the rebound response is specific to deprivation of nighttime quiescence rather than muscle fatigue.
“To distinguish between sleep deprivation and muscle fatigue, we performed the 6- or 12-hour mechanical stimulus experiments during the day (Figure 4B,E). We observed no significant difference between pre- and post-perturbation activity levels (Figure 4C,F), indicating that the rebound response is specific to deprivation of nighttime quiescence.”
Quote in the paperSubstantial claim- Rests on an assumption: Pulsing frequency is a valid measure of behavioral activity and quiescence in Cassiopea.
May combine two claims.
Findingown dataCassiopea jellyfish pulse less at night than during the day, with mean nighttime activity about 32% lower than daytime activity (night 781 ± 199 vs day 1155 ± 315 pulses per 20 min).
“We observed that Cassiopea pulse less at night than during the day (Figure 2; Data S1).”
- Also stated in: § RESULTS AND DISCUSSION · plain
“all showed a large decrease in mean activity (~32%) at night (781 ± 199 pulses/20 min, mean ± s.d.) compared to the day (1155 ± 315 pulses/20 min, mean ± s.d.; Figure 2C,E).”
Quote in the paperSubstantial claim- Rests on an assumption: Pulsing frequency is a valid measure of behavioral activity and quiescence in Cassiopea.
- Also stated in: § RESULTS AND DISCUSSION · plain
May combine two claims.
Findingown dataThe reduced nighttime responsiveness of Cassiopea is rapidly reversible: a second drop within 30 s decreased response latencies at night, and after the second drop day and night latencies were indistinguishable.
“During the night, there is a large decrease in the time to first pulse and time to reach the bottom, after the second drop when compared to the first drop (Figure 3D,E).”
- Also stated in: § RESULTS AND DISCUSSION · boosted
“During the day and night, the time to first pulse and time to bottom after the second drop were indistinguishable, demonstrating that after perturbation, animals have similar arousal levels during the day and night.”
- Also stated in: § RESULTS AND DISCUSSION · hedged
“These results indicate that Cassiopea have rapidly reversible reduced responsiveness to a stimulus during the night.”
Quote in the paperSubstantial claim- Rests on an assumption: Pulsing frequency is a valid measure of behavioral activity and quiescence in Cassiopea.
- Also stated in: § RESULTS AND DISCUSSION · boosted
May combine two claims.
Findingown dataDaytime feeding does not cause the day-night behavioral difference; no-feeding recordings were consistent with feeding recordings.
“To ensure that day feeding does not cause the day-night behavioral difference, we tracked the activity of 16 jellyfish over three consecutive days and nights without feeding and observed results consistent with those including feeding (Figure 2F,G; Figure S3D).”
Quote in the paperSubstantial claim- Rests on an assumption: Pulsing frequency is a valid measure of behavioral activity and quiescence in Cassiopea.
May combine two claims.
Findingown dataCassiopea activity decreased through the first 3-6 hours of night, with lowest activity 6-12 hours after the day-to-night transition.
“Jellyfish activity decreased throughout the first 3–6 hours of the night, with the lowest activity occurring 6–12 hours after the day to night transition.”
Quote in the paperSubstantial claim- Rests on an assumption: Pulsing frequency is a valid measure of behavioral activity and quiescence in Cassiopea.
Classified as probably not a substantial claim.
Findingown dataCassiopea prefer staying on solid surfaces.
“We observed in our nursery that Cassiopea prefer staying on solid surfaces as is found in nature.”
Quote in the paperSubstantial claimMay combine two claims.
Findingown dataThe 95th percentile of the night IPI frequency distribution is 13.9 s, whereas the day 95th percentile is 2.5 s.
“These pauses are seen as a tail in the IPI frequency distribution (Figure 2B: 95th percentile of night IPI frequency distribution (gray) is 13.9 s). Such long pauses are rarely seen during the day (Figure 2B: 95th percentile of day IPI frequency distribution (yellow) is 2.5 s).”
Quote in the paperSubstantial claim- Rests on an assumption: Pulsing frequency is a valid measure of behavioral activity and quiescence in Cassiopea.
May combine two claims.
Findingown dataThe decreased activity after perturbation is due to both decreased pulsing frequency and increased pause length.
“This period of decreased activity is due to both decreased pulsing frequency (increased mode of IPI-length) and increased pause length (increase in the IPI-length 95th percentile) (Figure S4B,C).”
Quote in the paperSubstantial claim- Rests on an assumption: Pulsing frequency is a valid measure of behavioral activity and quiescence in Cassiopea.
May combine two claims.
Findingown dataAt night, the inter-pulse interval is typically longer than during the day, due to a longer mode of the IPI distribution and more frequent pauses.
“During the night, the IPI is typically longer than during the day (Figure 2A,B; Data S1; Figure S3A). Two features contribute to this lengthening of the IPI: (1) the mode of the IPI distribution is longer at night than during the day, and (2) night pulsing is more often interrupted by pauses of variable length.”
Quote in the paperSubstantial claim- Rests on an assumption: Pulsing frequency is a valid measure of behavioral activity and quiescence in Cassiopea.
May combine two claims.
PremisecitationCassiopea rarely swim and usually remain stationary with their bell on a surface.
“They rarely swim and rather remain stationary with their bell on a surface, hence their name, the upside-down jellyfish (Figure 1B; Figure S1A; Movie S1; [25,26]).”
Quote in the paperSubstantial claim0 of 2 sources checked- cites [25] Jantzen, 2010, Enhanced pore water nutrient fluxes by the upside-down jellyfish Cassiopea sp. in a Red Sea coral reef, Mar. Ecol. Prog. Ser. source
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- cites [26] Holland, 2004, Global phylogeography of Cassiopea (Scyphozoa: Rhizostomeae): molecular evidence for cryptic species and multiple invasions of the Hawaiian Islands, Mar. Biol. source
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May combine two claims.
Interpretationown dataThe activity decrease after 12-hour nighttime perturbation was larger than after 6-hour perturbation, indicating that sleep rebound magnitude depends on the amount of sleep deprivation.
“The decrease in activity caused by the 12-hour perturbation was larger than that of the 6-hour perturbation, indicating that the amount of sleep rebound is dependent on the level of sleep deprivation.”
Quote in the paperSubstantial claimProbably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.
PremisecitationA fully functioning circadian system is not essential for sleep; animals with null mutations in circadian rhythm genes still sleep, though sleep timing is altered.
“A fully functioning circadian system is also not essential for sleep to occur; animals with null mutations of circadian rhythm genes still sleep, though sleep timing is altered [30].”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [30] Zimmerman, 2008, Conservation of sleep: insights from non-mammalian model systems, Trends Neurosci. source
Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.
Study design and conductown dataAt least two laboratory replicates were performed within the recording setup, which holds up to 8 jellyfish.
“For these experiments we performed at least two laboratory replicates within our recording setup, which is limited to 8 jellyfish.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S3 (load-bearing).
PremisecitationSleep has been hypothesized to have multiple functions, including synaptic homeostasis, neurotransmitter regulation, cellular damage repair, toxin removal, memory consolidation, and energy conservation.
“It has been hypothesized that sleep has multiple functions, including synaptic homeostasis, regulation of neurotransmitters, repair of cellular damage, removal of toxins, memory consolidation and energy conservation [7], although the ancestral role and selective advantage of sleep remains elusive.”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [7] Joiner, 2016, Unraveling the evolutionary determinants of sleep, Curr. Biol. source
Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.
PremisecitationIn many animals sleep is regulated by both homeostatic and circadian systems, but this is not always the case.
“In many animals sleep is regulated by both homeostatic and circadian systems [29], but this is not always the case [4–7,30].”
Quote in the paperNot a substantial claim0 of 6 sources checked- cites [29] Borbély, 1999, Sleep homeostasis and models of sleep regulation, J. Biol. Rhythms
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- cites [4] Hill, 2014, Cellular stress induces a protective sleep-like state in C. elegans, Curr. Biol. source
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- cites [5] Trojanowski, 2016, Call it worm sleep, Trends Neurosci. source
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- cites [6] Allada, 2008, Unearthing the phylogenetic roots of sleep, Curr. Biol. source
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- cites [7] Joiner, 2016, Unraveling the evolutionary determinants of sleep, Curr. Biol. source
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- cites [30] Zimmerman, 2008, Conservation of sleep: insights from non-mammalian model systems, Trends Neurosci. source
Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing).
Study design and conductnone givenNo specific randomization method was used.
“No specific method for randomization was used.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S3 (load-bearing).
Study design and conductnone givenExperimental animals were assigned to groups at random.
“Animals were randomly assigned to experimental groups.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.
PremisecitationC. elegans exhibits a developmentally regulated sleep state, and adult C. elegans show a non-circadian stress-induced sleep state.
“For instance, the nematode C. elegans exhibits a developmentally regulated sleep state, and adult C. elegans show a non-circadian stress-induced-sleep state [4,5,31].”
Quote in the paperNot a substantial claim0 of 3 sources checked- cites [4] Hill, 2014, Cellular stress induces a protective sleep-like state in C. elegans, Curr. Biol. source
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- cites [5] Trojanowski, 2016, Call it worm sleep, Trends Neurosci. source
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- cites [31] Nath, 2016, C. elegans stress-induced sleep emerges from the collective action of multiple neuropeptides, Curr. Biol. source
Not checked in the free run.
May combine two claims.
Study design and conductown dataLog-transformed time-to-first-pulse data were approximately Gaussian distributed with equal variance between groups, validating standard two-way ANOVA and unpaired t-tests on the transformed data.
“Tests of the log transformed time to first pulse data showed that the transformed data was approximately Gaussian distributed with equal variance between experimental groups, validating the use of standard two-way ANOVA and unpaired t-tests on the transformed data.”
Quote in the paperSubstantial claimProbably not a substantial claim: may not pass S3 (load-bearing).
Study design and conductnone givenNo statistical method was used to predetermine sample size.
“No statistical methods were used to predetermine sample size.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S3 (load-bearing).
Study design and conductnone givenExperimental groups compared statistically were tested for equal variance.
“Experimental groups that were statistically compared were tested for equal variance.”
Quote in the paperNot a substantial claimClassified as probably not a substantial claim. May combine two claims.
PremisecitationMelatonin promotes sleep in diurnal vertebrates, including zebrafish and humans, and induces quiescence in invertebrates.
“One such conserved molecule, melatonin [32], promotes sleep in diurnal vertebrates, including zebrafish [33] and humans [34], and induces quiescence in invertebrates [35].”
Quote in the paperSubstantial claim0 of 3 sources checked- cites [32] Peres, 2014, Developmental and light-entrained expression of melatonin and its relationship to the circadian clock in the sea anemone Nematostella vectensis, Evodevo source
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- cites [33] Zhdanova, 2001, Melatonin promotes sleep-like state in zebrafish, Brain Res. source
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- cites [34] Brzezinski, 2005, Effects of exogenous melatonin on sleep: a meta-analysis, Sleep Med. Rev. source
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Probably not a substantial claim: may not pass S3 (load-bearing).
Study design and conductown dataNormality tests indicated all datasets were approximately Gaussian distributed except the time-to-first-pulse arousal data.
“The normality tests showed that all datasets were approximately Gaussian distributed with the exception of the time to first pulse arousal data.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S3 (load-bearing).
Study design and conductnone givenInvestigators were not blinded to allocation during experiments or outcome assessment.
“Investigators were not blinded to allocation during experiments and outcome assessment.”
Quote in the paperNot a substantial claimClassified as probably not a substantial claim.
PremisecitationAt least 600 million years of evolution separate cnidarians from bilaterians.
“Though at least 600 million years of evolution separate cnidarians from bilaterians [9–16], many aspects of the nervous system are conserved, including neuropeptides and neurotransmitters [15–20].”
Quote in the paperSubstantial claim0 of 8 sources checked- cites [9] Dunn, 2014, Animal phylogeny and its evolutionary implications, Annu. Rev. Ecol. Evol. Syst. source
Not checked in the free run.
- cites [10] Arendt, 2016, From nerve net to nerve ring, nerve cord and brain--evolution of the nervous system, Nat. Rev. Neurosci. source
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- cites [11] Katsuki, 2013, Jellyfish nervous systems, Curr. Biol. source
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- cites [12] Erwin, 2002, The last common bilaterian ancestor, Development source
Not checked in the free run.
- cites [13] Hejnol, 2015, Neural nets, Curr. Biol. source
Not checked in the free run.
- cites [14] Kelava, 2015, Evolution of eumetazoan nervous systems: insights from cnidarians, Philos. Trans. R. Soc. Lond. B Biol. Sci. source
Not checked in the free run.
- cites [15] Bosch, 2017, Back to the basics: cnidarians start to fire, Trends Neurosci. source
Not checked in the free run.
- cites [16] Grimmelikhuijzen, 1995, The nervous systems of cnidarians, EXS
Not checked in the free run.
Classified as probably not a substantial claim.
PremisecitationPyrilamine, a histamine H1 receptor antagonist, induces sleep in vertebrates.
“Pyrilamine, a histamine H1 receptor antagonist that induces sleep in vertebrates [36], also induces concentration-dependent quiescence in Cassiopea (Figure S4F).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [36] Gandhi, 2015, Melatonin is required for the circadian regulation of sleep, Neuron source
Not checked in the free run.
The paper relies on this without support.
Pulsing frequency is a valid measure of behavioral activity and quiescence in Cassiopea.
“We define activity as the total number of pulses in the first 20 minutes of each hour.”
“To monitor jellyfish activity, pulsing information was extracted from the individual frames of each recording.”
The paper relies on this without support.
Laboratory-recorded pulsing behavior reflects the jellyfish's natural behavioral state.
“Cassiopea were reared in artificial seawater (ASW, Instant Ocean, 30–34 ppt) at pH 8.1–8.3, 26–28°C with a 12-hour day/night cycle.”
“Individual jellyfish were placed into 700 mL square clear plastic containers (cubbies), with white sand bottoms, in 35 L (10 gallon) glass tanks (Figure S1C–F).”
Claims other papers make about this one
Each is checked against this paper’s own text: does it say what the citing paper says it does?
Sleep has been identified in every animal that has been carefully studied.
“Sleep has been recognized in each animal carefully scrutinized.”
From The Biology of General Anesthesia from Paramecium to Primate (2019) · cites this paper as [29]
Behaviorally quiescent or sleep-like states may be as globally widespread as susceptibility to volatile anesthetics.
“behaviorally quiescent or sleep-like states may be as globally penetrant as susceptibility to volatile anesthetics [22–29].”
From The Biology of General Anesthesia from Paramecium to Primate (2019) · cites this paper as [29]
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Flagged first, then by citations
Research works printed but not linked: 1
- Folmer, 1994, DNA primers for amplification of mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates, Mol. Mar. Biol. Biotechnol., 3, 294
Cited by
Sorted by citations
Neuronal Mechanisms for Sleep/Wake Regulation and Modulatory Drive
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Mitochondrial origins of the pressure to sleep
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- Authors
- Nath, R. D., Bedbrook, C. N., Abrams, M. J., Basinger, T., Bois, J. S., Prober, D. A., Sternberg, P. W., Gradinaru, V., & Goentoro, L.
- Affiliations
- California Institute of Technology (US)
- Corresponding author
- Goentoro, Lea
- ORCID
- Venue
- Current Biology
- Publisher
- Elsevier BV
- Published
- 2017-10
- Volume
- 27
- Issue
- 19
- Pages
- 2984-2990.e3
- ISSN
- 0960-9822 (ISSN-L)
- PMC
- PMC5653286
- OpenAlex
- W2756734175
- Cited by
- 260
- References
- 43
- Citations by year
- 2026: 26 · 2025: 28 · 2024: 32 · 2023: 22 · 2022: 38 · 2021: 28 · 2020: 27 · 2019: 31 · 2018: 21 · 2017: 7
- FWCI
- 16.71
- Type
- journal article
- Language
- en
- Open access
- bronze
- doi.org
- Licence
- www.elsevier.com/tdm/userlicense/1.0/ (tdm)
- www.elsevier.com/legal/tdmrep-license (tdm)
- www.elsevier.com/open-access/userlicense/1.0/ (vor, 373 d)
- doi.org/10.15223/policy-017 (stm-asf)
- doi.org/10.15223/policy-037 (stm-asf)
- doi.org/10.15223/policy-012 (stm-asf)
- doi.org/10.15223/policy-029 (stm-asf)
- doi.org/10.15223/policy-004 (stm-asf)
- Locations
- Current Biology (publishedVersion)
- PubMed (publishedVersion)
- Europe PMC (PubMed Central) (acceptedVersion)
- PubMed Central (acceptedVersion)
- CaltechAUTHORS (California Institute of Technology) (submittedVersion)
- Funders
- NIH Director’s New Innovator Award/PECASE (IDP20D017782-01)
- James S. McDonnell Foundation for Complex Systems Science (220020365)
- NIMH under a Ruth L. Kirschstein National Research Service Award (F31MH102913)
- NINDS under a Ruth L. Kirschstein National Research Service Award (F31NS100519)
- National Science Foundation Graduate Research Fellowship (1144469)
- NIH (T32GM007616)
- Heritage Medical Research Institute
- HHMI (047-101)
- Topics
- Physical Sciences › Earth and Planetary Sciences › Paleontology › Marine Invertebrate Physiology and Ecology
- Life Sciences › Neuroscience › Cognitive Neuroscience › Sleep and Wakefulness Research
- Life Sciences › Neuroscience › Endocrine and Autonomic Systems › Circadian rhythm and melatonin
- Keywords
- sleep, Cassiopea, box jellyfish, Cnidaria, sleep homeostasis, sleep evolution, animal evolution
- MeSH terms
- Animals, Biological Evolution, Sleep, Scyphozoa
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