Networks of ultra-low-power nodes capable of sensing, computation, and wireless communication have applications in medicine, science, industrial automation, and security. Over the past few years, deployments of wireless sensor networks (WSNs) have utilized nodes based on off-the-shelf general purpose microcontrollers. Reducing power consumption requires the development of Systemon-chip (SoC) implementations that must provide both energy efficiency and adequate performance to meet the demands of the long deployment lifetimes and bursts of computation that characterize WSN applications. This work takes a holistic approach and, thus, studies all layers of the design space, from the applications and architecture, to process technology and circuits. This paper introduces the emerging application space of wireless sensor networks and describes the motivation and need for a custom system architecture. The proposed design fully embraces the accelerator-based computing paradigm, including acceleration for the network layer (routing) and application layer (data filtering). Moreover, the architecture can disable the accelerators via VDD-gating to minimize leakage current during the long idle times common to WSN applications. We have implemented a system architecture for wireless sensor network nodes in 130nm CMOS. It operates at 550 mV and 12.5 MHz. Our system uses 100x less power when idle than a traditional microcontroller, and 10-600x less energy when active.