He received his Bachelor of Science degree with Honors in Botany from the University of Iowa, Iowa City, Iowa. His Honors thesis examined whether species of the genus Atrichum had sun and shade leaf forms and this initiated his life-long interest in both bryophytes and photosynthesis. He received his PhD in 1999, from the Botany Department at the University of Wisconsin-Madison, where he pursued research in photosynthesis and isoprene emission. He then went on to a post-doctoral fellowship at the Australian National University from 2000-2002 where he worked on Rubisco kinetics and CO2 concentrating mechanisms in hornworts and algae before taking up is current position.