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Thus, one or three eyespots were equally effective compared to a pair of eyespots, as long as the total area of the eyespots was conserved, suggesting that resemblance to a pair of vertebrate eyes is inconsequential.
They further found that markings with centres displaced inwards, making them seemingly three-dimensional and eye-like, were attacked no differently than non-eye-like markings without centre displacement .
They went on to conduct another study where they compared the effectiveness of eyespots with a black pupil surrounded by a yellow ring, which most closely resembles vertebrate eyes, with eyespots having other colour combinations.
Their study indicated that several other non-eye-like colour combinations were as effective as black and yellow  found that symmetry in size, shape and colour patterns is crucial for averting predatory attacks, thus indirectly supporting the eye-mimicry hypothesis.
Two broad hypotheses explain how eyespots may be effective against predation.
Large, conspicuous eyespots are considered to be intimidating for the predator thereby decreasing the chances of attack – the “Intimidation Hypothesis”, whereas smaller eyespots closer to the wing margin are thought to attract attention toward themselves thus deflecting predatory attacks away from the more vital parts of the prey – the “Deflection Hypothesis”  demonstrating the role of eyespots in reducing predation paved the way for further intensive studies establishing that eyespots can indeed intimidate predators.
Although the intimidating effect of eyespots has been shown in multiple studies, there is still a lack of a deeper understanding of the mechanistic basis of “why” they intimidate .
Lepidopteran insects have been especially popular for investigations into eyespot evolution.
Major strides have been taken toward understanding the genetics and developmental processes involved in eyespot formation in butterflies, particularly through studies on laboratory populations of ].
Two competing hypotheses seek to explain the cause of intimidation, one suggesting ‘eye-mimicry’ and the other their ‘conspicuousness’ as the reason.
There is an on-going debate about which of these better explains the effectiveness of eyespots against predation.