In a perfect system, no A-a gradient would exist: oxygen would diffuse and equalize across the capillary membrane, and the pressures in the arterial system and alveoli would be equal (resulting in an A-a gradient of zero). However even though the partial pressure of oxygen is about equilibrated between the pulmonary capillaries and the alveolar gas, this equilibrium is not maintained as blood travels further through pulmonary circulation. As a rule, is always higher than by at least 5–10 mmHg, even in a healthy person with normal ventilation and perfusion. This gradient exists due to both physiological right-to-left shunting and a physiological V/Q mismatch caused by gravity-dependent differences in perfusion to various zones of the lungs. The bronchial vessels deliver nutrients and oxygen to certain lung tissues, and some of this spent, deoxygenated venous blood drains into the highly oxygenated pulmonary veins, causing a right-to-left shunt. Further, the effects of gravity alter the flow of both blood and air through various heights of the lung. In the upright lung, both perfusion and ventilation are greatest at the base, but the gradient of perfusion is steeper than that of ventilation so V/Q ratio is higher at the apex than at the base. This means that blood flowing through capillaries at the base of the lung is not fully oxygenated.