At early stages of heart development, the first and second heart fields are a continuum of lateral head mesoderm-derived, cardiogenic cells.

Pioneering work in the chicken established that the initial development of the heart consists of two stages: the quick assembly of a beating heart, followed by the recruitment of cells from adjacent tissues to deliver the mature in-and outflow tract. Cells to build the primitive heart were dubbed the first heart field (FHF) cells, cells to be recruited later the second heart field (SHF) cells. The current view is that these cells represent distinct, maybe even pre-determined lineages. However, it is still unclear where exactly FHF and SHF are located at different stages of development, and whether there is a sharp boundary or rather an overlap between the two. It is also unclear whether both FHF cells and SHF cells originate from the lateral head mesoderm (LHM), whether the paraxial head mesoderm (PHM) contributes to the SHF, and where the LHM-PHM boundary might be. To investigate this problem, we exploited the size, ease of access and exquisite anatomy of the chicken embryo and used traditional strategies as well as newly developed transgenic lines to trace the location of cardiogenic fields and boundaries from the time the first heart-markers are expressed to the time SHF cell recruitment ceases. Our work shows that both FHF and SHF stem from the LHM. We also found that FHF and SHF lack a distinct anatomical boundary. Rather, FHF and SHF are a continuum, and the recruitment of cells into the heart is a chance event depending on morphogenetic movements, the position of cells within the moving tissues, the separation of the somatic and splanchnic LHM, and the separation of the heart from the splanchnic subpharyngeal mesoderm during heart-looping. Reconciling our and previous studies we propose that first and second heart field precursors are specified but not determined, thus relying on morphogenetic processes and local environments to realise their cardiogenic potential.