Theacrine is a small alkaoid molecule which is very similar to caffeine in both structure and effects. The available evidence suggests it is actually synthesized from caffeine in certain plants.
The differences in caffeine and theacrine are particularly interesting, most significantly that research suggests a tolerance to the effects of theacrine (as Teacrine®) forms at a far slower pace than it does to caffeine.  In addition, theacrine has pain reducing and anti-stress properties that caffeine does not. 
The available evidence suggests the boosts in mental energy, focus, and motivation from theacrine are provided through a similar mechanism as in caffeine, through both the adenosinergic and dopaminergic pathways.
Like caffeine, Theacrine's primary mechanism of action likely involves it's effects on the brain's adenosine receptors.  Adenosine is a chemical which can be found in nearly all human cells, but plays a particularly important role in the brain. Every second you're awake, your brain cells are constantly firing and as they do so they release adensoine as a byproduct.
When released in the brain, adenosine functions as an inhibitory neurotransmitter. This means that when it binds to adenosine receptors it slows down brain activity and acts as a central nervous system depressant. When adenosine binds with a receptor, it also causes blood vessels to expand, allowing more oxygen to reach the brain during sleep.
Typically as adenosine binds to its receptors and slows down nerve activity this manifests in drowiness and fatigue. After someone has been awake all day, adenosine has continued to accumulate. Once levels are high enough, sleepiness sets in.
Caffeine molecules have a very similar shape as adensoine molecules. The shapes of each molecule are so similar in fact, that when caffeine is consumed it can bind to adenosine receptors, preventing adenosine from binding with them. This causes a reduction in fatigue and increases in mental energy. Studies on theacrine have shown very similar changes in adenosine signalling when theacrine is administered as when caffeine is administered, so it is believed theacrine functions much in the same way. 
Research on Theacrine has also found that it may influence dopaminergic signalling in a similar way as caffeine influences adenosine signalling.  Dopamine is an excitory neurotransmitter (meaning it causes stimulation), and plays key roles in motor control, arousal, motivation, cognitive functioning, and learning.
Studies on theacrine have shown that concentrated doses activate the dopaminergic receptors D1 and D2, causing an increase in average dopamine levels while the theacrine is active.  Elevated dopamine levels are commonly associated with improved mood, motivation, focus, and mental clarity. Research has shown theacrine (as Teacrine®) to produce many of these benefits, especially improvements in motivation and focus.  Theacrine's ability to raise dopamine levels is the most likely cause of these benefits.
In animal studies theacrine was found to possess a number of benefits that caffeine does not, pointing to further differences in the mechanisms of action for the two supplements. One study found Theacrine to have pain reducing effects in mice comparable to those of the drug indomethacin. 
Another study found that theacrine supplementation lowered the severity of stress induced liver damage in mice, suggesting that it has anti-stress properties. 
It is not yet known by what mechanism Theacrine produces it's anti stress and pain reducing effects, but it's influence on the dopaminergic and adenosinergic pathways is unlikely, suggesting it has additional mechanisms of action that are not yet well understood.
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