They can act from a distance as much as a few millimeters, as well as act in short range. Netrins can also act as a repellant. Unique among chemoattractants is that the function of netrin has been conserved among a variety of species across 600 million years. Like netrin, ephrin can function as both an attractant and repellant. Ephrins primarily play a role in setting a gradient along the anterior-poster axis for the guidance of developing retinal axons. Semaphorins, which were first identified on specific axons in the grasshopper CNS, function primarily as short-range inhibitory cues that steer pioneer axons away from less ideal regions. Receptor complexes for semaphorins include neuropilins and plexins. The Slit-Robo cell signaling pathway plays an important role in guiding pioneer axons, especially pioneer longitudinal axons. These axons, which function to connect major parts of the CNS, are mainly present during embryonic development. The Slit family mainly functions as a repellent towards longitudinal axons, guiding them away from the ventral midline. The loss of Slit in "Drosophila" caused the presence of longitudinal axons in the midline. In conjunction with the Robo receptor, Slit signaling played a role in determining tract positions parallel to the midline for longitudinal axons to follow during development. The loss of either Slit or Robo caused dysfunction in the development of longitudinal pioneer neurons in the midbrain and hindbrain of "Drosophila". Furthermore, it has been shown that Robo plays a diversified role in pioneer axon guidance in different areas of the brain during embryonic development. Primarily, Robo 1 is crucial towards pioneer longitudinal axon guidance in the ventral tract, while Robo 2 is important in the dorsal tract. The signaling associated with the receptor Notch, as well as non-canonical Notch/Abl signaling, have been shown to play a role in the development of longitudinal pioneer neurons in the "Drosophila" ventral nerve cord. The Notch receptor has been shown to interact with interface glia to form a path that longitudinal pioneer neurons can follow. Notch/Abl signaling in the pioneer neurons increases the motility of the growth cones of longitudinal pioneer axons while stimulating filopodia development. It has also been noted that Notch signaling is also important in the migration of neurons in the mammalian cortex. The directed growth of axons depends on structure at the end of the tip of a growing axon referred to as a growth cone.