20Comparative gene expression studies are invaluable for predicting how existing genetic 21 pathways may be modified or redeployed to produce novel and variable phenotypes. Fruits are 22 ecologically important organs because of their impact on plant fitness and seed dispersal, 42 siliques within the Brassicaceae, facilitating future studies on seed shattering in important 43 Brassicaceous crops and pernicious weeds.44 45 3 46 62Arabidopsis, especially amongst close relatives e.g., the loss of dehiscence in many species 63 across the Brassicaceae (1).
65Brassicaceae fruits vary markedly in shape, structure, and size (1,8). Their variation in 66 dehiscence is a focal point for research because it fundamentally changes fruit structure, 67 subsequently affecting dispersal and diversification (9). A prerequisite for exploring how 68 differences in fruit morphology are achieved across the Brassicaceae is familiarity with both the 69 fruit structure and underlying genetic pathways in Arabidopsis (10,11). Arabidopsis fruits, 102 results in indehiscent fruits in Arabidopsis: SHP1/2, SPT,. Overexpression 103 of FUL or NO TRANSMITTING TRACT (NTT) also results in indehiscent fruits (28,29); FUL 104 overexpression completely suppresses SHP1/2, resulting in reduced lignification in the enb layer 105 and reduced valve margin formation; overexpression of NTT phenocopies the ful mutation 106 resulting in valve margin specific genes being expressed throughout valve. In summary, a 107 modification of many components in this pathway results in a loss of dehiscence. Because 108 indehiscence is observed in at least 20 different lineages across the family, it is likely that this 109 phenotype evolved via multiple modifications to this pathway (30). As such, there is no singular 110 alteration to the fruit patterning pathway implicated in this shift for all tribes. 111 112 To date, little is known about the genetic basis of indehiscence in the Brassicaceae, although it is 113 currently being bridged by studies in taxa with varying indehiscent morphologies. Recently, a114 study demonstrated a deviation in expression of eight key genes between pod shatter sensitive 115 species and shatter resistant species of Brassica and Sinapis (2). In Lepidium, there has been an 116 evolutionary shift from dehiscence to indehiscence, e.g., valve margin genes that are conserved 6 117 between the dehiscent L. campestre and Arabidopsis have been lost in the indehiscent L.118 apellianum (31,32). Upregulation in upstream regulator AP2 has been suggested as a factor in 119 this indehiscence (32).120 121 A notable morphological adaptation is the evolution of a complex fruit type known as 122 heteroarthrocarpy, which is only found in some members of the tribe Brassiceae (30,33,34). This 123 modified silique is defined by the presence of a variably abscising central joint, an indehiscent 124 distal region, and a variably dehiscent proximal region (Fig 2). As such, this novel morphology 125 offers an opportunity to investigate fruit variation beyond shifts from ...