Over the following half-century, most physicians and researchers among the Pima Indians avoided the conclusion that a more sedentary lifestyle was the decisive factor in the degradation of health on communities. Instead, many drew a correlation with the increase in processed starches and high sugar beverages among reservation Pima communities, in contrast to those who lived away from those regions that came to rely on federally-sanctioned processed foods. The health of these Native American communities, then, was impacted by new federal food and land policies; just as earlier colonial interventions impacted their nutritional lives in problematic ways.Among other resources for such a conclusion and/or postscript to their course, students could use Annual Reports of the Commissioner of Indians Affairs. The reports provide evidence from Indian Territory between the 1820s and the 1930s, describing nutritional changes, diseases, perceived health problems, and ecological and agricultural developments. Students could also assess Indian and Pioneer Papers, which detail thousands of interviews of residents of Oklahoma in the 1930s. Many of those residents were elderly and recalled nutritional, agricultural, and health developments that took place over the previous century. Students – and future researchers – might even consult advertisements from “Indian Territory” newspapers. Their proliferation of cures, tonics, and “snake-oil” medicines for digestive problems after the Civil War era might provide clues to problematic dietary changes among Native Americans; changes that resulted in part from disruptive state and federal policies that were contiguous with challenging interventions that took place during the earlier colonial eras.Into the most recent era, indeed, reservation communities have been more susceptible to certain problems that are associated with the modern western diet,vertical farm as broadly defined during the second half of the twentieth century – particularly its reliance on refined carbohydrates in packaged and processed goods.
From the 1960s to the 1970s, for example, medical researchers began to define a clear correlation between the appearance of obesity and Type II diabetes in Native American communities and their adoption of processed foods in federal welfare programs that targeted newly formed reservations. Reports of Type II diabetes among Oklahoma communities increased after their move into federally-mandated reservations through the 1940s. A review of literature from 1832 to 1939 by the epidemiologist Kelly West found no reports of diabetes among the Kiowa, Comanche or Apache communities living in the region .Similarly, diabetes seems to have been much less prevalent among Pima Indian populations who lived away from reservations in the American Southwest before 1940. By the turn of the twenty-first century, however, around half of all Pima Indians have been reported to suffer from diabetes.Following their analysis of early contact history, students might consider these more recent phenomena in light of modern scientific literature on metabolic syndrome. According to federal statistics from the present day, moreover, Native American communities continue to suffer from diabetes, cirrhosis, influenza, pneumonia, and perinatal and early infancy diseases at greater rates than the general American population.Yet as the USDA nutritional guidelines for the Food Distribution Program on Indian Reservations program demonstrate, Native American populations will continue to receive food welfare in the form of packaged and processed starches, seed oils, and low fat animal sources – foods that have potentially contributed to increasing metabolic syndrome in the US population more generally.As they try to understand possible correlations between the two scholarly fields of history and nutritional science, therefore, students and researchers might begin to think about problematic nutritional interventions and paradigms outside the Native American community, among the American public more generally.
The study of early-American history and the problematic European intervention in Native American nutritional life should inform their future careers in fields such as public health, scientific research, nutritional science, and medicine. Approximately five to seven million years ago, nectar spurs arose in the ancestor of Aquilegia , which then diversified into ∼ 70 species in two major clades: one distributed across Eurasia, and the other primarily distributed across North America. Spur morphology varies substantially across the genus, ranging in length from ∼ 1-16 cm, and varying in other morphological characteristics such as the degree of curvature and color. These petal characteristics contribute to pollinator specificity, which plays an important role in reproductive isolation between taxa . For example, species that occur in both Eurasia and North America have the ancestral morphology of the genus — shorter, curved spurs, usually ranging in color from blue to purple — and are typically pollinated by bees. In North America, convergent evolution has lead to several lineages with red and yellow flowers, and straight, medium-length nectar spurs that are primarily pollinated by hummingbirds. Multiple North American lineages that generally lack floral anthocyanins and are yellow or white have evolved long nectar spurs and are pollinated by hawk moths. In addition to these three pollination syndromes, one species native to montane regions of central China, A. ecalcarata, has secondarily lost nectar spurs and is primarily pollinated by syrphid flies. Given the role that nectar spurs have played in the diversification of Aquilegia, understanding the genetic and developmental basis of how they form is key to understanding both the initial evolution of this three dimensional structure as well as the generation of subsequent modifications that serve as adaptations to different pollinators.Phase I broadly comprises the mitotic phase of petal development in which localized cell-divisions establish the spur cup. Initially, cell divisions are dispersed throughout the petal primordium, but they cease in a wave that begins at the margins of the petal, moving basipetally and causing divisions to become concentrated in the nascent spur.
Mitotic activity is maintained in the developing spur until the spur reaches 5-9 mm in length, but it is progressively restricted toward the spur tip, where the nectary develops. As cells begin to differentiate in Phase II, anisotropic cell expansion becomes a major contributor to spur shape,nft vertical farming particularly length. Differences in spur length across Aquilegia species have been primarily attributed to differences in cell length, rather than cell number, although the basis of spur curvature and other aspects of shape have yet to be studied in detail. In an effort to understand the genetic basis of nectar spur development in Aquilegia, a previous study examined gene expression differences between the petal blade and spur at several early developmental stages in the horticultural variety A. coerulea ‘Origami’. This study ruled out a role for type I KNOX genes, which maintain cellular indeterminacy in meristematic tissue, indicating that the prolonged mitotic activity in the spur is not meristematic in nature. However, a number of other genes whose homologs play a role in regulating the transition between cell proliferation and differentiation in the petals of Arabidopsis thaliana were highly differentially expressed between the petal blade and spur. For example, the TEOSINTE BRANCHED/CYCLOIDEA/PCFgene, AqTCP4, whose homolog in A. thaliana, TCP4, controls petal size by repressing cell-division, was more highly expressed in petal blades where mitotic activity first ceases. In contrast, a homolog of an A. thaliana GRF-INTERACTING FACTORgene that controls petal size by promoting cell-division, AqGIF1/AN3, was up-regulated in the Aquilegia spur where mitotic activity is maintained. The most highly differentially expressed gene identified in this study is a member of the STYLISHgene family, whose homologs in A. thaliana are best known for their functions in carpel, rather than petal, development. Subsequent analyses of this gene, AqSTY1, and two additional Aquilegia homologs in the STY gene family, AqSTY2 and Aq LATERAL ROOT PRIMORDIUM , revealed that in addition to a conserved role in carpel development, these genes are critical to nectary development. Although the AqSTY-like genes do not appear to function in the earliest phases of nectar spur development, these results highlight the utility of gene expression studies to identify novel candidates involved in unique roles of Aquilegia petal development. The previous gene expression and functional analyses were conducted on a single Aquilegia cultivar, and did not address questions regarding the conservation and divergence of gene expression patterns during early petal development across the diverse morphologies in the genus Aquilegia. Here we conduct transcriptomic analyses of Phase I petals at five different developmental stages from four different Aquilegia species, A. ecalcarata, A. sibirica, A. formosa, and A. chrysantha, representing a diverse set of pollination morphologies and spanning the Aquilegia phylogeny . This broad sampling allowed us to take a comparative approach, combining differential expression and weighted gene correlation network analyses to explore both commonalities in the genetic basis of petal development across the genus Aquilegia as well as taxon-specific differences.
To that end, a set of genes commonly DE between the three spurred species in our study, A. sibirica, A. formosa, and A. chrysantha, and the spurless species, A. ecalcarata was compared to the set of DE genes between the petal blade and spur previously conducted in A. coerulea ‘Origami’. This comparison revealed only 35 genes that are either more highly expressed in both A. ecalcarata and the petal blade or in the spurred taxa and the petal spur. In addition to these unbiased analyses, we explored the expression of Aquilegia homologs of genes known to regulate the transition between cell division and differentiation in A. thaliana petals in order determine the potential for broad functional conservation as well as any potential role in Aquilegia spur development.Petals were dissected from floral buds spanning Phase I of development from each of four species of Aquilegia – A. ecalcarata, A. sibirica, A. formosa, and A. chrysantha. Although it can be challenging to determine homologous developmental stages across taxa, a combination of petal morphology and stamen development was used to group samples across the four species into five developmental stages , DS1-DS5, with petals from at least three flowers collected for each species and stage. Representative samples of each species at each of the five developmental stages are presented in Fig. 2. The first petal stage assessed was collected from buds approximately equivalent to floral stage 8. At this point, the morphology of petals from all four species examined was quite similar. Petals were approximately 0.5 mm wide and, although they exhibited slight concave adaxial curvature, the spur had yet to initiate. Notably, this is the first time that a pre-initiation stage petal has been sampled for RNA analysis in Aquilegia. The second petal stage examined came from floral buds at approximately stage 9. At this stage, the spur had begun to form in the spurred taxa and was approximately 0.5 mm long while petals were approximately 1 mm wide in the blade region. Even at this early stage of nectar spur development, morphological differences between the taxa were already detectable, with the developing spur of A. formosa being wider than those of A. sibirica and A. chrysantha . These differences in the spurred taxa continued through the third petal stage sampled where developing spurs continued to elongate but largely maintained their shape as established in DS2. By the fourth petal stage sampled , additional morphological differences became apparent. For example, although spurs were only between 1 and 2 mm long at this stage, curvature in the A. sibirica petal was already being established. Some of the spurs at this stage began to appear more bulbous at the tip of the spur, indicating the incipient development of the nectary. Although A. ecalcarata does not develop a spur or a nectary, at DS4 a small pocket is visible below the petal attachment point. In addition to spur differences, several subtle differences in petal blade morphology across the taxa were apparent, including the shape of the blade apex and its relative length compared to the spur. The developmental gap between the fourth and fifth petal stages collected was larger than the gaps between the other stages. At the fifth time point collected , the spurs of the spurred taxa ranged in length from 4-7 mm and morphological differences were accentuated. A. sibirica spurs exhibited more extreme curvature, A. formosa spurs were broad through most of the length of the spur, narrowing just before the nectary, and A. chrysantha spurs were long and narrow. In general, this stage should represent the transition out of phase I into phase II, where cell elongation is the dominant factor affecting spur length.We started by comparing changes in the transcriptional profiles of petals between serial developmental stages for each species. For A. ecalcarata, A. sibirica, and A. chrysantha, the differences in gene transcription were minimal between each of the first four stages, whereas there was a larger jump between DS4 and DS5 .