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This is a very serious problem, which causes at least 60% performance loss.

Using VisualVM, I found that glBindVertexArray took up a lot of CPU time.

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I think this is caused by frequent OpenGL calls after glBindVertexArray, such as glBindBuffer, glEnableVertexAttribArray, glVertexAttribPointer. 

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VAO only needs to bind its attributes once, and then it does not need to bind every time it is drawn.

Finally, a large number of drawing calls will also cause performance loss. Try merging vertex buffers into one.