The detection of life in extreme environments and the amazing capacity of microbes to obtain energy from the environment has led to the hypothesis that heat, not water or energy sources, is the final barrier that constrains life on Earth. As such, life is expected to extend far into Earth's crust, potentially inhabiting an environment as extensive as the world's oceans [1]. This chapter aims to summarize what is currently known about life in the oceanic crust, which contains all three domains of life and is a vastly underexplored habitat. This is not the first summary of this topic, and additional content can be found in numerous review and primary research papers [2-16], book chapters [17,18], and books [19][20][21].The discovery of hyperthermophilic archaea and recurring geochemical evidence of crustal alteration led to the first hypotheses of a subsurface biosphere in the Earth's crust [22]. Initial suggestions of this subsurface biosphere came from experiments at seafloor hydrothermal vents, in which elevated DNA levels in hydrothermal plumes were measured that could not be explained by simple seawater entrainment, implied that microbes were likely to live in underlying sediments and rocks. Around this time, advances were also made with experiments showing that microbe could and should grow on the Earth's crust, furthering the concept of a subsurface crustal biosphere [23][24][25]. Despite the difficulty of sample retrieval, and low and heterogeneous biomass on crust samples, experimental evidence has now confirmed the existence of crustal life, consisting of a vast microbial community including viruses (reviewed in [5, 6, 13, 14, 26]).The early evidence for a crustal biosphere inevitably led to lines of inquiry intended to address questions concerning its extent, global biomass and transport and distribution. The crustal biosphere extends over a wide temperature range, with evidence for biogenic alteration in the temperature range from 15-80°C [1]. Yet, most of the subseafloor oceanic crust is at least in theory habitable, based on measured and modeled isotherms, which show a conservative 120°C isotherm to extend thousands of meters into the crust, which is below the predicted (150°C; [22]) and experimentally measured temperature maxima of life (122°C [27]). The volume of this potentially habitable zone in ocean crust equals that of the modern ocean and exceeds that within continental crust by nearly a factor of 2 [1]. Over geologic time, the creation and cooling of new crust has increased the volume that is potentially habitable. Based on comparBrought to you by | Stockholms Universitet Authenticated Download Date | 8/25/15 7:03 AM