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Home / A Review of Visual Perception Mechanisms That Regulate Rapid Adaptive Camouflage in Cuttlefish

A Review of Visual Perception Mechanisms That Regulate Rapid Adaptive Camouflage in Cuttlefish

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Introduction

Cuttlefish are adept at rapidly altering their body patterning and skin texture for camouflage (Hanlon and Messenger, 1988, 1996; Shohet et al., 2006; Kelman et al., 2007; Mäthger et al., 2007; Allen et al., 2009; Zylinski et al., 2009a, b, c; Barbosa et al., 2012). This expeditious expression of cover-up torso patterns is a visually driven behavior (Holmes, 1940; Hanlon and Messenger, 1988; Marshall and Messenger, 1996) that enables cuttlefish to adjust to homogeneous surroundings (e.g., sand plains) and also complex habitats, such as coral reefs, kelp forests, and temperate rock reefs, with relative freedom from detection or recognition by their many visual predators (Boletzky, 1983; Hanlon, 2007; Hanlon et al., 2011).

Cuttlefish take several camouflage tactics to choose from when trying to avoid visual predation (Hanlon et al., 2009). They primarily deceive predators through background matching, i.e., resembling the background to hinder detection; disruptive patterning, i.e., obscuring edges, shape, and body outline to impede detection or possibly recognition; or masquerade, which is resembling an inanimate or uninteresting object to avoid recognition or detection (e.k., Cott, 1940; Hanlon and Messenger, 1988; Hanlon et al., 2009; Stevens and Merilaita, 2009). The basic camouflage body patterns used by cuttlefish tin be grouped into three categories: Compatible, Mottle, and Disruptive (Hanlon, 2007). Each torso pattern is made upwards of low-cal and dark splotches that range along a continuum (Hanlon et al., 2009). The Uniform design is composed of minor splotches, the Mottle body pattern features medium-sized splotches, and the Confusing torso pattern consists of large splotches forth with bars and stripes (Hanlon and Messenger, 1988).

Cuttlefish are benthic animals that dwell primarily on the seafloor, where they utilize visual information from the surrounding substrate and objects in their immediate vicinity to conform their advent for cover-up. This information includes horizontal cues, such equally sand and gravel, vertical facades (e.g., wall, rock face), too equally discrete, 3D objects (e.g., rocks, algae clumps) (Hanlon and Messenger, 1996; Barbosa et al., 2008a; Buresch et al., 2011). Contempo studies have highlighted the importance of vertical features, such as 3D objects and 2D representations of 3D objects, in eliciting the expression of cuttlefish camouflage body patterning (Barbosa et al., 2008a; Buresch et al., 2011; Ulmer et al., 2013). In ane set of experiments, cuttlefish preferentially masqueraded as high-contrast, 3D objects that occupied only a minor percent of the visual field in their environment (Buresch et al., 2011). In another experiment, vertical features alone had a stronger influence on body patterning than horizontal, benthic cues (Ulmer et al., 2013).

Although a growing torso of literature is get-go to unravel those visual cues that arm-twist dissimilar body patterns for camouflage (e.thou., contrast, aspect ratio, shape, substrate intensity, spatial stage, edges, and design size; Hanlon and Messenger, 1996; Chiao and Hanlon, 2001a, b; Chiao et al., 2005, 2009, 2010, 2013; Barbosa et al., 2007, 2008a, b; Shohet et al., 2007; Kelman et al., 2008; Hanlon et al., 2009; Zylinski et al., 2009a), the distance at which visual cues are relevant for camouflage has not been studied empirically. This series of experiments addressed two specific questions: (1) How far away from the animal are visual cues relevant for camouflage? and (2) How exercise 2D and 3D visual stimuli—and their orientation and altitude—influence torso blueprint choice for groundwork matching or masquerade?

Materials and Methods

Animals

European cuttlefish (Sepia officinalis Linnaeus, 1758) were hatched, reared, and maintained in the Marine Resources Heart facility of the Marine Biological Laboratory in Woods Pigsty, Massachusetts. Xx animals (boilerplate pall length (ML) = 5.8 cm, SD = 0.five cm; average White Square area = 2.25 cm2, SD = 0.4 cm2) were used for these experiments. (For a total description of the White Square, come across Chiao and Hanlon, 2001a.) We also measured each animate being'due south body length (BL; boilerplate BL = 8.2 cm, SD = 0.vii cm), and used these values as a distance measure for the experimental substrates and objects.

Experimental setup

Experiments were conducted in a circular, 42-cm bore tank with flow-through seawater with a depth of 10 cm. The tank was located within a blackness tent to prevent disturbances during the experimental trials. A circular, 37 cm diameter, LED white light source (High Effulgence LED strip lights; Environmental Lights, San Diego, CA) was placed directly above the arena to reduce the result of shadows. To maintain a consistent distance of presented stimuli, cuttlefish were placed in a clear, plexiglass cylinder (14.v cm diameter, 14.5 cm acme) located in the center of the experimental tank. Animals could rotate within this cylinder, merely could not swim a significant altitude. Cuttlefish commonly settled within this cylinder and showed a stable torso pattern within several minutes.

Animals were tested individually, and both animal and stimulus orders were randomized. Animals were given 15–60 min to acclimate to the tank; an animal was considered acclimated when it showed a stable body pattern with little or no motility. Cuttlefish were observed on a TV monitor outside the tent, and images were taken remotely (Canon Insubordinate XS, Catechism United statesA., Inc., Melville, NY) later the animate being had settled. I image was taken for each cuttlefish on each substrate per 3D object combination.

As was shown earlier, cuttlefish use iii basic body pattern types for camouflage: Compatible, Mottle, and Disruptive, with variations on each design type (Hanlon et al., 2009; for a description of each body pattern type, see Hanlon and Messenger, 1988). In the current experiments, we used large-scale, high-contrast checkerboards, which are known to arm-twist the Disruptive body design, to investigate the body pattern response of South. officinalis, because the visual cues used to evoke this pattern have been studied extensively (Chiao and Hanlon, 2001a, b; Mäthger et al., 2006, 2007; Barbosa et al., 2007, 2008a; Chiao et al., 2007, 2009; Kelman et al., 2007; Zylinski et al., 2009a, b). In these experiments, cuttlefish may use Disruptive body patterning to background lucifer the blackness and white checkerboard on the substrate and/or the wall, or to masquerade as a 3D object.

All stimuli were positioned in the same identify for each animal. Bogus substrates and objects were made using uniform gray reckoner printouts (RGB = 142; 50% gray) designed to elicit a Uniform trunk pattern. Large black (RGB = 255) and white (RGB = 0), high-contrast checkerboard squares were fashioned to elicit a Disruptive body pattern (checkerboard square size = 2.25 cm2, equal to 100% of the animals' boilerplate White Foursquare area). Substrates were computer-generated and laminated to exist waterproof. A single 3D object, 6.0 cm in diameter (approximately equal to 1 ML) and 6.0 cm loftier, was constructed using the same black and white checkerboard squares used for the substrate flooring and wall.

Control experiments

Two command experiments were performed, during which (1) a uniform, fifty% grayness substrate floor and 25-cm-bore arena wall were used to arm-twist Compatible patterning; and (2) a checkerboard substrate floor and loonshit wall were used to elicit Disruptive patterning. For both control experiments, the walls were presented directly against the exterior of the plexiglass cylinder (at 0BL).

Experiment i: Horizontal floor (background matching)

(See Fig. 1A for experimental design.) Cuttlefish were presented with loftier-contrast checkerboard squares located on the 2nd floor at 3 distances: a) directly underneath the cuttlefish in a 1BL-diameter circle; b) directly exterior the 0BL diameter of the plexiglass cylinder, with a 50% gray floor located underneath the cuttlefish; and c) a band effectually the cuttlefish at 1BL (viii.0 cm from plexiglass), with a 50% gray flooring located directly beneath the cuttlefish, extending to the 1BL distance. For Experiment 1, a l% grayness wall was placed along the edge of the experimental loonshit.

Figure 1.
Figure 1.

Experimental setup based on the known response of cuttlefish to produce Confusing trunk patterns in response to checkerboards of the appropriate size and dissimilarity. Cuttlefish were placed inside a articulate plexiglass cylinder and presented with either: (A) a blackness and white checkerboard floor placed a) directly beneath the cuttlefish, b) at 0 body lengths (BL) away from the cuttlefish, or c) ane BL away from the cuttlefish; (B) a vertical black and white checkerboard wall placed a) at 0 BL abroad from the cuttlefish, or b) 1 BL away from the cuttlefish; or (C) a blackness and white, 3D cylinder placed at a) 0 BL away from the cuttlefish, b) i BL abroad from the cuttlefish, or c) 2 BL away from the cuttlefish.

Experiment ii: Vertical wall (background matching)

(Meet Fig. 1B for experimental design.) Cuttlefish were presented with high-contrast checks located on the second vertical wall at two distances: a) at 0BL, correct against the clear cylinder; and b) at 1BL (8.0 cm from the plexiglass). For Experiment 2, a 50% gray floor covered the entire experimental arena.

Experiment 3: 3D object (masquerade)

(Come across Fig. 1C for experimental design.) A high-contrast, 3D object was moved to three distances from the cuttlefish: a) 0BL, i.e., right against the plexiglass cylinder; b) 1BL (eight.0 cm from the plexiglass); and c) 2BL (16.0 cm from the plexiglass). For Experiment iii, a 50% gray wall was placed along the edge of the experimental arena, and a 50% grey floor covered the entire experimental arena.

Paradigm assay

We used a MATLAB R2010a-generated image analysis plan (The Mathworks, Inc., Natick, MA), developed by C. Chiao, C. Chubb, and 50. Siemann, as an automated method for characterizing and discriminating between cuttlefish body patterns (for more detail, see Chiao et al., 2009). This programme performs a fast Fourier transform of each prototype and analyzes the paradigm in unlike spatial frequency bands, assigning an free energy level to each of the half-dozen bands. The three cuttlefish body patterns: Uniform, Mottle, and Disruptive, differ in spatial calibration (or granularity), and can exist distinguished by the singled-out shape of their granularity spectra. In addition, this computer program uses landmarks (assigned past the user) on the cuttlefish trunk to locate 11 Disruptive body blueprint components within the image. A "Disruptive design score," based on the relative dissimilarity of the pixels in each component, is then generated for each image.

Statistical differences in Disruptive pattern scores by substrate were analyzed using a 1-mode repeated measures ANOVA in the MATLAB statistics toolbox. Pairwise comparisons between substrates were made using a Tukey-Kramer test in the multcompare function in MATLAB.

Results

Command experiments

A ane-way repeated measures ANOVA showed a meaning difference in body patterning between substrates (F = 22.99; P = 2.76 eastward-25). Cuttlefish showed a Uniform body pattern on the 50% gray control and a Confusing body pattern on the checkerboard command (Fig. two). Pairwise comparisons from a Tukey-Kramer test showed a pregnant difference betwixt the Disruptive torso design score on the grayness control substrate versus the checkerboard command [greyness command: M (mean) = 0.fifty, checkerboard control: M = 6.4; P < 0.05].

Figure 2.
Figure 2.

Trunk patterning responses to each experimental background (see Fig. one). The Disruptive score is determined by the relative dissimilarity between pixels within each paradigm. Box plots represent the range of Disruptive body design scores, the line depicts the median, and fault bars are S.D. Two asterisks (**) indicate which substrates differed significantly from the grayness command (*). The cuttlefish drawings illustrate the body blueprint elicited by each visual background. BL, body length; Checker, checkerboard.

Experiment one: Horizontal floor

Cuttlefish responded to loftier-contrast checkerboard squares located on the substrate directly beneath them with weak Disruptive body patterning (i.due east., White Foursquare only), only did not respond with Confusing coloration to checkerboard squares located across their immediate 0BL annulus (Fig. 2). Pairwise comparisons showed a significant divergence between the Disruptive body design score on the gray control substrate and the checkerboard floor placed straight beneath the cuttlefish (grayness control: Thousand = 0.50, checkerboard flooring: Thou = 2.43; P < 0.05), only not betwixt the body design response on the gray control and the response to the floor when it was 0BL and 1BL abroad (0BL: Thou = 1.21, 1BL = 0.86; P > 0.05).

Experiment two: Vertical wall

Animals responded to high-contrast, vertical checkerboard squares located at 0BL with a Disruptive trunk design. The Disruptive response decreased markedly at 1BL (Fig. two); in some cuttlefish, simply the White Square was expressed and others were Uniform. Pairwise comparisons showed a meaning deviation between the Disruptive body pattern score on the grey command and the checkerboard wall located at 0BL (checkerboard wall 0BL: K = 4.71; P < 0.05), simply not between the body pattern responses to the grayness control and the checkerboard wall at 1BL (checkerboard wall 1BL: M = i.61; P > 0.05).

Experiment iii: 3D object

Animals responded with weak Disruptive torso patterning (i.e., White Square but) to a high-contrast, 3D object placed at 0BL and 1BL (Fig. 2). When the checkerboard object was placed at 2BL, some animals connected to respond with weak Confusing body patterning, while others became Uniform (Fig. 2). Pairwise comparisons showed a significant divergence between the Disruptive body pattern score on the gray command and the checkerboard object located at 0BL and 1BL (checkerboard object 0BL: M = 2.49; checkerboard object 1BL: M = two.21; P < 0.05), but not between the torso design response to the gray command and the checkerboard object at 2BL (checkerboard object 2BL: M = 1.75; P > 0.05).

Give-and-take

In these experiments, nosotros set out to address how far abroad visual stimuli are relevant to cuttlefish camouflage body patterning, and to assess whether there was a divergence in their response to 2D horizontal and vertical stimuli. In general, cuttlefish responded with Disruptive body pattern elements only to visual stimuli that were within one body length of distance, whether the stimuli were presented horizontally or vertically. Disruptive body pattern response macerated quickly as stimuli were presented at distances greater than 1 body length. In addition, the cuttlefish Confusing body design response was strongest in response to both 2D and 3D vertical stimuli than to benthic, horizontal stimuli. This result was not unexpected, since piece of work from our laboratory has shown that vertical stimuli have more influence over cuttlefish body patterning than horizontal stimuli (Barbosa et al., 2008a; Ulmer et al., 2013).

Cuttlefish depend on their unique camouflage abilities for survival. Since their marine environment is oftentimes complex and heterogeneous in nature, they must exist able chop-chop to alter their cover-up trunk patterning to blend into their environment, or to masquerade as nearby objects (e.chiliad., rocks, algae). There are ii possible explanations for their use of visual stimuli that are very close by. Outset, it would seem near effective, in deceiving predator vision, for their camouflage to blend with immediately adjacent backgrounds. Second, as a benthic species, cuttlefish probably experience low-visibility conditions on a regular basis. Since cuttlefish have large and sensitive eyes (Groeger et al., 2005), and can cover-up themselves in extremely low-low-cal conditions (Allen et al., 2010b; Buresch et al., 2015), information technology is unlikely that their vision is affected much past turbid water. However, the vision of many potential fish predators may be affected by water turbidity (Utne-Palm, 2002). Cuttlefish thus may choose to resemble but nearby objects and substrates, because information technology is probable they are detected only when a predator is near.

Surprisingly, cuttlefish responded to 3D objects farther away than they did to 2nd substrates. This finding suggests that the visual sampling rules for masquerade may differ from those used in background matching. While cuttlefish responded merely to 2D substrates that were very close by (i.e., 0BL from the brute), they continued to reply to 3D objects when these were placed 2BL from the fauna. These results suggest that cuttlefish are more than likely to utilize masquerade than background matching as a cover-up tactic if a loftier-dissimilarity, 3D object is inside a few body lengths away. One reason may be that patterns on the substrate become too distorted for the cuttlefish to recognize when observing them at a well-nigh grazing incidence. It is non possible to prove definitively that the camouflage tactic used by the cuttlefish in our experiments was masquerade; such proof would require viewing the cuttlefish from the perspective of a predator (Stevens and Merilaita, 2009; Skelhorn et al., 2010). However, the behavior was similar to that of cuttlefish in the wild––using body posture and texture plus chromatic trunk patterning to resemble inanimate objects in their surroundings (Hanlon et al., 2009). This finding fits the recent definition of masquerade: "looking similar an inedible or inanimate object" (Skelhorn et al., 2010, 2011; Skelhorn and Ruxton, 2011). Other cephalopods use this type of masquerade for cover-up, selectively sampling just a few visual features from their surroundings for body patterning (Hanlon et al., 1999). Ii octopus species, Octopus cyanea and Octopus vulgaris, base some of their torso patterns on features of nearby objects rather than on their entire field of view (Josef et al., 2012).

Another interesting issue of our experiments was that cuttlefish appeared to be able to perceive the distance of the 3D objects. Camouflage body patterning is by and large scale-dependent: when the background scale changes, cuttlefish deploy an appropriately scaled body pattern. That is, large-scale elements elicit Disruptive torso patterns; medium-scale elements bring out Mottled trunk patterns; and small-scale elements, Uniform body patterns (Barbosa et al., 2008b; Chiao et al., 2009). The cuttlefish in this series of experiments appeared to identify the scale of the patterns on the objects presented to them, equally noted by their use of a Disruptive body design in response to the large-scale checkerboard object. In some cases, they used Disruptive body patterning fifty-fifty when the object was upward to 2BL abroad. At this distance, the change in perspective would take fabricated the checkerboard squares appear to be 25% of their original size at 0BL. This checkerboard square size (25% of the cuttlefish's own White Square area) would commonly arm-twist a Mottle torso pattern if presented directly below or abreast a cuttlefish (checkerboard squares 12%-forty% of an animal's White Foursquare size elicit a Mottle body pattern; Barbosa et al., 2008b). However, cuttlefish responded with a Disruptive torso design. It is unclear if the apparent ability of cuttlefish to appraise the altitude of the 3D objects in this experiment––and therefore the actual size of the checkerboard squares––was due to the 3D nature of the object itself or to some other mechanism in the cuttlefish eye.

Recent studies suggest that cuttlefish possess depth perception capability (Josef et al., 2014). Yet since their optics are laterally placed, they must be able to conform in some way the depth necessary to recognize that objects are located at a distance. Ii theories take been offered: accommodation is achieved past motion of the lens perpendicular to the axis of the eye (Schaeffel et al., 1999); and, in sure circumstances, the "W" shape of the cuttlefish pupil may aid in depth perception (Mäthger et al., 2013). Regardless, the cuttlefish in our experiments appeared to judge the distance of the 3D objects that were presented to them. In addition, at that place is evidence that cuttlefish perceive 3D substrates differently from 2D substrates, and that visual depth may really increment the strength of some Disruptive components (Kelman et al., 2008), including the White Foursquare––the Confusing component elicited by the objects in this set of experiments. It is possible that the visual depth of the 3D objects may be the ultimate cause of the difference in cuttlefish response to the 2d versus 3D cues that we noted in our experiments.

In the wild, cuttlefish live in a wide variety of natural environments that incorporate many different objects with visual depth (e.1000., rocks, algae, and coral; Hanlon and Messenger, 1988; Hanlon et al., 2011). They have often been observed to camouflage themselves as these objects, even when they are at a altitude of 5 body lengths away (R. Hanlon, pers. obs.). On the other hand, cuttlefish more often than not merely utilize background matching to cover-up to substrates that are immediately beneath them or in which they are partially cached (Hanlon and Messenger 1988; Hanlon et al., 1999, 2009; Allen et al., 2010a). These experiments, along with many field observations, highlight the importance of masquerade as a choice of cover-up tactic in this species.

Acknowledgments

Nosotros thank the animal care staff in the Marine Resources Center for help with weekend care of our cuttlefish colony. Liese Siemann and Justine Allen provided valuable insight and discussion for this study. Charlie Chubb provided help with statistics. This piece of work was funded by DARPA/DSO grant no. W15P7T-13-D-CT04.

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Source: https://www.journals.uchicago.edu/doi/full/10.1086/BBLv229n2p160

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