Plant nitrogen nutrition is an essential and energy-costly component of terrestrial food chains. Understanding nitrate sensing in plants can lead to improved crop yields and nutrient use efficiency, directly impacting food security and agricultural sustainability. Herein, we review and present a comprehensive framework for understanding nitrate sensing in plants, integrating molecular, genetic, and physiological aspects. We begin by detailing the primary nitrate response and nitrate starvation response, which are central to the plant's ability to sense and respond to nitrate availability. We then explore the intricate interactions between nitrate signaling and other nutritional pathways such as those for carbon, phosphorus, potassium, and sulfur assimilation and reactive oxygen species (ROS) handling, and how it unfolds in long-distance systemic communication between roots and shoots. Finally, evolutionary insights are provided by comparing nitrate-sensing mechanisms across different plant species as well as Bacteria, Archaea, Chlorophyta, Charophyta (algae), and Fungi, revealing how these mechanisms may have evolved in diverse ecological niches. This review not only provides a framework to project our present and future understanding of plant nitrate and nitrogen nutrition but also offers potential strategies for improving nutrient use efficiency in crops through genetic and biotechnological interventions.