Not freshly washed shirt = smells like hasn't been washed

I have a shirt that has been hanging in a closet which is well-ventilated (it consists of a hanging rail with no doors). It was washed about a year ago and has not been worn since its last wash, i.e. a year ago. It’s been hanging on the same rail as my other clothes that I wear regularly. But unlike my other garments, this one has an odour (which could be best described as ‘not clean and fresh’). This would generally be attributed to poor ventilation, but the ventilation on my door-less clothes rail is much better than the conventional wardrobe with doors and yet this still seems to happen with shirts that haven’t been worn in a while (not those that were washed in the last couple of months).

So WHAT is the cause of the ‘unwashed’ smell in shirts that have indeed been washed, albeit they were washed a long time ago? Is it something to do with bacteria and/or dust mite eating your dead skin cells on the shirt, and if so is there a health risk to wearing the shirt despite the smell (if you can put up with the smell, that is)?