When do field mice sleep
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Need Help Managing Pests? Chem-Free offers both effective, low-impact pest control options and preventative measures to help avoid future infestations. We examined the performance of our automatic method of state detection. On six arbitrarily chosen 4-h long segments, the agreement of automatic detection against experienced user detection was The major disagreement was in the detection of very brief states typically microarousals which were often not detected not considered by an experienced observer.
Figure 2. Distribution of sleep—wake states in young and older mice over 24 h. A Distribution of wake, SWS, and REM sleep recorded over 23 consecutive days with 1-h resolution in individual mice black lines and their average red line.
Each line represents averaged values of one animal over several days of recordings, the red line is an average from the example mice shown in panel A. Shaded area indicates the dark phase. Printed values are estimated marginal means computed in six independent mixed linear models one for each state of vigilance and phase of light—dark cycle with age and hour factors.
Printed values are estimated marginal means from independent mixed linear models. Note that older mice show more SWS than young mice over a day, but this difference occurs mainly during the dark phase. We analyzed the automatically detected state distribution for 99 days from eight young mice and 67 days from nine older mice. Out of at least 3 weeks of recording per animal, we typically analyzed 12—14 days from each animal, but up to 23 days, starting from the second week after electrode implantation and connection to recording cables.
Some days were skipped because of occasional technical difficulties. The same animal typically demonstrated very similar sleep—wake pattern across days Figure 2A except occasional long wake states during the day, likely due to unusual environmental conditions atypical noise in animal facilities.
The older mice had a more variable sleep—wake behavior. Post hoc tests in the light phase showed that old animals had less REM at the beginning of the cycle. Significance was not reached at any time points for post hoc of age effects on wake during light phase.
Finally, main effect of age on SWS duration was non-significant in the light phase. Overall, these results suggest that differences in sleep-wake behavior of young vs. Most states in mice were short. As it can be seen from Figures 3A—C , at the beginning of the dark phase of cycle, when the sleep pressure is low, the animal continuously switched between SWS and waking states. The SWS was often interrupted by brief waking episodes characterized by an activated LFP in all channels and increased muscle tone Figure 3B as well as during wake, we often observed episodes of reduced muscle tone and high power of LFP activities in the low-frequency bands, characteristic for SWS Figure 3C.
Using 5 s resolution for state detection, very brief muscle twitches typically less than a second , even accompanied with activated LFPs were not detected as wake not shown. During REM sleep, the LFP was typically activated and the muscle tone was mostly absent with the exception of occasional twitches.
Overall bimodal distribution of SWS suggests that SWS episodes lasting 10 s and less represent some separate states and can be considered as microsleep states Figure 3D.
For the wake, the short episodes had likely two dominant durations with a clear maximum around 7 s and less obvious around 20 s Figure 3D. It is possible that these two types of electrophysiological microarousal states represent some different behavioral states.
Figure 3. Fragmentation of sleep and wake states. A One-hour long LFP recordings at the beginning of the dark phase — h in a 1-year old mouse. B,C Expended fragments from this recording demonstrating the presence of very short wake episodes during SWS [activated LFP and increased muscle tone, B ] and sleep episodes during wake [slow-wave activity and reduced muscle tone, C ]. D Share of all episodes from all investigated animals and days of wake, SWS, and REM sleep of different duration with 1 s resolution for 24 h, for light phase and dark phase as indicated for young and older mice.
These distributions show that older mice have larger number of short-lasting states, in particular during the dark phase. During SWS, cortical neurons have a bi-stable behavior alternating between active and silent states and during REM sleep or wake, the membrane potential has a unimodal distribution Steriade et al.
Therefore, during wake or REM sleep, the ratio of activities below and above 4 Hz delta range is nearly 1, but during SWS the activity in the delta range power is six times higher than the faster frequencies power, reflecting the bi-stable behavior of cortical neurons Mukovski et al. In our experiments, the overall delta power distribution was bimodal, with low values corresponding to waking state and REM sleep and high values corresponding to SWS Figure 1D. The origin of delta activity during wake or REM sleep is unclear; therefore, we further analyzed the delta power dynamics only during SWS.
Previous experiments on cats demonstrated that intracellular sleep slow wave activity had different amplitude in different cortical areas Chauvette et al. Therefore, we calculated the mean hourly delta power 0. As expected from sleep homeostasis process Borbely, ; Borbely et al. Moreover, post hoc tests showed that older mice have a significantly higher delta power in the frontal cortex, while there was no difference in the somatosensory anterior and posterior cortex for both light and dark phases.
As it is clearly seen in Figure 4A , in older mice, the overall delta power was the highest in the frontal cortex, lower in anterior somatosensory, and even lower in posterior somatosensory cortices, while in young mice the delta power was the lowest in the frontal cortex and it was higher in both anterior and posterior somatosensory cortex which is the opposite of what is seen in human Carrier et al.
Figure 4. Area-specific daily SWS delta power dynamics is different in young and older mice. Printed values are estimated marginal means computed in two independent mixed linear models one per light or dark phase using factors of age, electrode, and time of the day.
B Mean SWS delta power in frontal cortex red , somatosensory anterior cortex green , and somatosensory posterior cortex black for 24 h left panel , light phase middle panel , and dark phase right panel for young bright colors and older light colors mice. The differences in SWS delta power in different ages and areas are likely mediated either by the number of waves or by their properties. Therefore, using a neural network approach Bukhtiyarova et al.
The differences were not significant for slow wave duration or amplitude. These results suggest that a higher incidence of slow waves was likely a factor of the increase in the delta power observed in the frontal cortex of older mice. Figure 5. Slow wave features in young and older mice. Slow waves are detected using a neural network approach and detected slow waves are depicted by black traces. B Expanded segment from A as indicated.
D Duration of detected slow waves, E Amplitude of detected slow waves. It is very common that the detection of states of vigilance in rodents is done based on just one or two electrodes. Using EEG recordings in mice, it was recently shown that spontaneous sleep Fernandez et al. Combined with our data on area-specific differences in delta power, these results suggest that various areas can have different propensities for sleep and wake states.
Therefore, we investigated LFP activities in 14 different cortical locations together with neck muscle EMG in intermediate age mice 6 months old, Figure 6. The overall idea of these experiments was to identify the given state of vigilance based on each LFP electrode separately and muscle activities, and then to compare the coincidence of state detection. Based on a formal criteria of state detection Figure 1 we found that a the transition between states does not occur simultaneously and even in closely located areas, the delays of state transition could take up to 20 s compare the first and second green traces, Figure 6A ; b while REM sleep high intensity theta rhythm and low muscle tone can be detected in a large number of cortical areas, some channels display slow-wave activities at the same time Figure 6B.
Using formal criteria of state identification, we often observed simultaneously the presence of two and sometimes three different states in different parts of neocortex Supplementary Movie S1.
Therefore, electrographic activities corresponding to SWS, REM sleep, or wake can co-occur in different cortical areas. Figure 6. Area-specific distribution of electrographic brain states. Signals from electrodes are color coded and the location of electrodes is indicated in the inserted drawing.
B A short segment that was overall qualified as REM sleep, but two fronto-laterally located electrodes show clear slow-wave activity. C h distribution of states detected based on muscle activity and each electrode individually. In the light phase, 3-months old and 1-year-old mice spent similar time in sleep and wake states except that 1-year-old mice spent significantly less time in REM sleep at the beginning of each period.
Similarly, 1-year-old mice spent less time in REM sleep at the end of the dark phase of the cycle. Generally, in the dark phase, older mice spent globally less time in wake and more time in SWS. The increase in the total sleep duration was mainly due to an increase in the number of short less than a minute sleep episodes during the dark phase.
The LFP delta power had remarkable regional specificity. Older mice showed larger LFP delta power in the frontal cortex compared to young mice, which can be explained by a tendency for an increase in slow wave density. We also investigated regional specificity of sleep—wake electrographic activities and found that using formal criteria, posterior regions of the cortex show more REM sleep activities high theta power during low muscle tone , while somatosensory cortex displays more often SWS patterns.
In agreement with multiple previous investigations, our study shows that overall, mice spent more time sleeping during the light part of the light—dark cycle and more time in waking state during the dark phase of the cycle Tobler et al. However, the formal definition of microstates is unclear.
However, recent advances in automatic methods of state detection demonstrated that the conventional states can be much more fragmented than it was previously assumed Koch et al. It is known that brain states in animals undergo rapid changes, which are associated with marked changes in neuronal activities in different structures Bezdudnaya et al. Human studies indicate that ms is the minimal time to mediate conscious perception Libet et al.
An essential condition for a conscious state to be generated is spatio-temporally coordinated neuronal firing Konorski, The presence of EEG slow wave activities is a key factor for the loss of consciousness Purdon et al.
Sleep or anesthesia-induced slow waves in cats and those measured intracellularly are mediated by neuronal hyperpolarization and silence that lasts — ms Timofeev et al. Therefore, the active states during sleep and anesthesia last typically longer than ms and the animals are still unconscious.
Thus, the minimal duration of an arousal should be longer than ms and likely longer than 1 s in order to be considered as a state.
On the contrary, the presence of local slow waves in some spatially restricted parts of the cortex alters behavioral responses Vyazovskiy et al. This suggests that they represent isolated states, which likely can be called microarousal or microsleep states. Similar activities exceeding this time likely represent either short or long states, not microstates. Because the delta power varied not only between cycles, but also between the beginning and the end of the cycle, and we used the same threshold throughout each phase light or dark , we could artificially increase the detection of short states.
However, there are no reasons to believe that the method would affect more the results of one group in particular; therefore, our observation of a higher number of short-lasting states wake or SWS in older mice compared to young mice is very solid. The state duration in mice living in natural environment could be different. For example, it is unlikely that mice foraging in the forest at the beginning of the dark phase of the sleep—wake cycle will have brief sleep episodes.
The anterior somatosensory electrode in our study was located in the barrel field and we did not observe age-specific differences in the sleep delta power in young vs. This suggests that there are very little age-dependent changes in the most important sensory system of mice. While no differences were found in the posterior somatosensory cortex trunk area , there was an increase in delta power and a trend for an increased slow wave density in frontal cortex of older mice.
Increase in delta power in frontal cortex in older mice are opposed to what is seen in humans Carrier et al. Mechanistic aspects of such differences are unclear and will require further studies. It was demonstrated in human that a reduction of cortical thickness was responsible for an age-dependent reduction in slow wave activities Dube et al.
There is no difference in frontal cortical thickness between 1- and 3. This might explain the relative stability of delta power in the barrel cortex in our experiments. Mice seemed to be a very attractive model to study aging. By the age of 3 months, they become mature adults, and at the age 10—15 months they belong to middle-aged group and after 18 months they are considered as being old Flurkey et al.
The aging in mice occurs substantially faster than in other laboratory animals and if the model is good, it would give multiple practical advances to study aging. During aging there are major alterations in human sleep. We took several appropriate parameters of human sleep affected by aging Mander et al.
Thus, while overall time spent in sleep is different between species, both show a similar increase of sleep during their active phase light for humans, dark for mice.
Different from human, in our mice experiments, the overall slow-wave activity did not undergo systematic changes. We found an increase with aging in the delta power and a trend for an increased slow wave density in frontal cortex, but stable delta power in the posterior part of somatosensory cortex. Although effects of aging in humans are more prominent in NREM sleep, reductions in REM sleep are also reported in healthy older participants Carrier et al. One aspect of our study investigated the spatial distribution of formally defined states over the cortical surface.
We found that electrodes implanted in frontal cortex would identify more of wake, in motor and somatosensory cortex more SWS and in visual and retrosplenial cortex more REM sleep Figure 6. REM sleep-related theta activity was more often seen in posterior parts of the dorsal cortical surface and slow-wave activity during otherwise REM sleep was often recorded over somatosensory and motor cortical areas Figure 6. This supports a recent discovery of REM sleep-dependent slow wave activity in mice Funk et al.
The observed dominance of wake, SWS, and REM sleep regions in our study largely overlaps with the anatomical localization of cortical clusters belonging to the default mode network identified in awake mice Vanni et al. Two types of slow-wave activities during REM sleep were also found in human Bernardi et al. Frontal—central slow waves, which authors believe are analogous with PGO waves, had increased gamma activity during slow waves Bernardi et al. It is well-demonstrated that sleep slow waves in both human and animals are associated with a reduction of gamma activities Mukovski et al.
These mice are prey to cats, dogs, bears, wolves, snakes, owls and rabbits. Nocturnal in nature, they are cautious and venture outside only after having fully assessed their surroundings. Mice use their coats as camouflage, blending in with rocks or dried leaves. Despite this, these mice are often consumed by their predators.
Most field mice do not survive to their second year. Females are able to become pregnant every month. Baby field mice are born blind, bald and deaf. However, within several weeks, they will have reached maturity and begin mating. Call Residential Commercial. Brown, black or white. What Orkin Does Orkin technicians are trained to help manage field mice and other rodents. Call us or. Get Your Quote.
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