Patterns of systemic stress during the agricultural transition in prehistoric Japan.код для вставкиСкачать
AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 142:112–124 (2010) Patterns of Systemic Stress During the Agricultural Transition in Prehistoric Japan Daniel H. Temple* Department of Anthropology, University of Missouri, Columbia, MO 65211-1440 KEY WORDS enamel hypoplasia; cribra orbitalia; Jomon; Yayoi; bioarchaeology ABSTRACT This study documents and interprets systemic stress during the agricultural transition in prehistoric Japan using linear enamel hypoplasia (LEH) defects and cribra orbitalia (CO) lesions. Middle to Final Jomon cultures (5000–2300 BP) from Honshu Island represent the foraging samples, while Yayoi cultures (2500–1700 BP) represent the early agricultural samples. Jomon foragers from eastern Japan had broad-based, intensive economies. Jomon foragers from western Japan had a greater focus on seasonally available, nutritionally poor resources, while Yayoi people were descendents of migrants from the East Asian continent and introduced wet rice economies to Japan. This study tests the hypotheses that wet rice economies will be associated with a lower prevalence of teeth/individuals affected by LEH defects in western Japan, while few differences in the prevalence of teeth/individuals with LEH defects will be observed between eastern Jomon people and Yayoi farmers. It is further predicted that similar CO prevalence will be observed between Jomon and Yayoi people given environmental similarities. Signiﬁcantly greater frequencies of teeth affected by LEH defects are observed among western Jomon compared to Yayoi people. The prevalence of teeth with LEH defects is slightly elevated among eastern Jomon foragers compared to Yayoi agriculturalists. Significant differences in CO prevalence are not observed. Systemic stress prevalence in western Japan likely declined following wet-rice agriculture because this crop provided a predictable, renewable resource base. Systemic stress prevalence was similar between eastern Jomon and Yayoi people because both groups practiced intensive subsistence strategies. Similar CO prevalence reﬂects infectious diseases associated with living conditions. Am J Phys Anthropol 142:112–124, 2010. V 2009 Wiley-Liss, Inc. This study documents and interprets systemic stress patterning during the agricultural transition in prehistoric Japan using linear enamel hypoplasia (LEH) defects and cribra orbitalia (CO) lesions. LEH defects and CO lesions were utilized because these nonspeciﬁc indicators of stress represent independent reactions to environmental stimuli and provide a broad perspective on health outcomes between behaviorally and ecologically diverse people (Huss-Ashmore et al., 1982; Goodman et al., 1984; Larsen, 1997). LEH defects are horizontal pits or grooves observable on tooth surfaces caused by disrupted enamel production during infancy and early childhood (Goodman and Rose, 1991). When enamel production is disrupted, shortened enamel prisms are deposited on the dentine by ameloblasts (enamel producing cells) (Hillson, 1996). Disruption of enamel production is associated with a variety of conditions including malnutrition, acute and chronic infection, trauma, and hereditary anomalies (Goodman and Rose, 1991). There is general consensus, based on observations among living communities and individuals from known environments, that the prevalence of teeth/ individuals affected by LEH defects reﬂects systemic stress levels (Goodman et al., 1991; May et al., 1993; Zhou and Corruccini, 1998). CO lesions occur when blood producing bone marrow in the cranium expands, increasing red blood cell production (Stuart-Macadam, 1985). Marrow expansion through the cortex of the orbital roof produces sieve-like lesions. CO is associated with a variety of anemic conditions, rickets, scurvy, and inﬂammation (Wapler et al., 2004; Walker et al., 2009). Many of these nutritional deﬁciencies are associated with parasite exposure and diarrheal disease (Walker, 1985, 1986; Walker et al., 2009). The agricultural transition represents one of the most pivotal phases of biocultural evolution for modern humans (Larsen, 1987, 1995, 2002). Mutual reliance between humans and plants characterizes this behavioral shift and suggests humans became dietarily dependent on agricultural products (Rindos, 1984). Such relationships differ from foraging or food procurement economies where domestication of plants (plant reliance on humans) often occurs in the absence of human reliance on cultigens (Harris, 1989; Smith, 2000). Agricultural economies formed the basis for expansions in population density and urbanization. Bioarchaeological research has questioned the extent to which modern humans beneﬁted from the transition to food producing economies (Cohen and Armelagos, 1984; Larsen, 1987, 1995, 2002, 2006; Cohen and CraneKramer, 2007). Percentages of teeth and individuals with LEH defects increased signiﬁcantly in response to dietary stress and infectious disease levels in the Western Hemisphere and Southwest Asia following the transition to intensive agriculture (Cook, 1980, 1981, 1984; C 2009 V WILEY-LISS, INC. C Grant sponsor: Wenner Gren Foundation for Anthropological Research. *Correspondence to: Daniel H. Temple, Department of Anthropology, University of Missouri-Columbia, Columbia, MO 65211-1440. E-mail: firstname.lastname@example.org Received 5 February 2009; accepted 3 September 2009 DOI 10.1002/ajpa.21208 Published online 1 December 2009 in Wiley InterScience (www.interscience.wiley.com). 113 JOMON AND YAYOI STRESS TABLE 1. Jomon and Yayoi sample data Site Kitamura Ota Tsukumo Yosekura Hobi Inariyama Nakazuma Yoshigo Doigahama Kanenokumac Koura Nagaoka Date 5000–3000 5000–4000 3000–2300 4000–3000 3000–2300 3000–2300 4000–3000 3400–2400 2500–1900 2500–1900 1900–1700 2100–1900 Group BP BP BP BP BP BP BP BP BP BP BP BP b Inland Jomon Western Jomon Western Jomon Western Jomon Eastern Jomon Eastern Jomon Eastern Jomon Eastern Jomon Yayoi Yayoi Yayoi Yayoi Curation Na Nagano Kyoto University Kyoto University Tokyo University Tokyo University Kyoto University Kyoto University Kyoto University Kyushu University Kyushu University Kyushu University Kyushu University 93 42 67 59 66 34 103 181 143 68 53 20 a Approximate numbers of individuals available for study. The number of teeth and orbital vaults available for each sample differs from these numbers. b Inland Jomon diet was likely similar to those from western regions (Akazawa, 1986; Minagawa and Akazawa, 1992). c A skeleton from this sample was dated to 2350 BP using AMS 14C methods with correction for the marine reservoir effect (Tanaka et al., 2005). Kennedy, 1984; Rose et al., 1991; Lukacs, 1992; Larsen et al., 2002; and others). CO prevalence increased among maritime foragers and following the transition to agriculture in many regions, reﬂecting the interaction of infectious disease loads with nutritional homeostasis (El-Najjar, 1976; Walker, 1985, 1986; Walker et al., 2009). The biological impact of wet rice dependent economies in Southeast Asia differs from that observed in other regions. Increases in LEH defects and CO prevalence are not reported in Thailand or Vietnam (Pietrusewsky and Douglas, 2002a,b; Oxenham, 2004; Domett and Tayles, 2007; Douglas and Pietrusewsky, 2007). Continued reliance on broad-based economies and stasis in frequencies of infectious diseases are likely responsible for these trends in Thailand (Pietrusewsky and Douglas, 2002a,b; Douglas and Pietrusewsky, 2007; Domett and Tayles, 2007), though increases in the prevalence and variety of infectious diseases are reported in regions such as Vietnam (Oxenham et al., 2005). These ﬁndings suggest that the transition to wet rice economies may not have produced the types of nutritional stressors associated with intensive agriculture in other regions of the globe, though chronic infection related to population aggregation and migration is noted (Larsen, 2002, 2006). In Japan, Jomon subsistence economies were broadbased, regionally variable, and terminated with the introduction of wet rice agriculture during the Yayoi period (2500–1700 BP) (Akazawa, 1981, 1982, 1986; Akazawa and Maeyama, 1986; Imamura, 1996a,b; Habu, 2004). Geographic distributions of tools indicate that Jomon foragers from eastern Japan had more sophisticated systems for maritime and terrestrial mammal food procurement than those from western and inland regions (Akazawa, 1986; Akazawa and Maeyama, 1986). Eastern Jomon economies were supported by a forest– estuary ecosystem that included coastal embayments, long-spanning coastlines, and diluvial uplands (Akazawa, 1986). Forest–estuary ecosystems likely provided year-round access to foods such as ﬁnﬁsh and mollusks, terrestrial mammals, and plant products suggesting that calorically dense foods were available for much of the year (Akazawa, 1986, 1999; Minagawa and Akazawa, 1992). In contrast, western and inland Jomon sites were situated in forest–freshwater ecosystems that exist as transitional zones between rivers, lakes, and forests (Aka- zawa, 1999). Tools related to plant procurement discriminate sites from these regions (Akazawa, 1986; Akazawa and Maeyama, 1986). Plant availability in this region was subject to seasonal ﬂuctuations and may not have been available at certain times of the year (Akazawa, 1986; Hudson, 1999). Nevertheless, recent stable isotope analysis of human skeletal remains suggests signiﬁcant consumption of marine ﬁnﬁsh by inland/western compared to eastern Jomon people (Minagawa, 2001; Yoneda et al., 2004; Kusaka, personal communications). Finﬁsh consumed by these groups was, however, seasonal in availability (Kobayashi, 2005). Fallback foods exploited by western Jomon people were nutritionally poor, particularly when compared with those consumed by eastern Jomon communities (Akazawa, 1986). Fallback foods are deﬁned as resources consumed during periods when primary/preferred foods are unavailable (Marlowe and Berbesque, 2009). These fallback foods included acorns, leafy greens, roots, tubers, and walnuts in western Japan (see: Akazawa, 1986 Table 1) and terrestrial mammals combined with seasonal roots/tubers in eastern Japan (Akazawa, 1999). Greater frequencies of teeth and individuals with LEH defects are observed among Jomon people from western and inland sites compared to eastern regions (Shigehara, 1994; Temple, 2007a). These studies suggest western and inland Jomon people experienced greater systemic stress due to narrow food availability and nutritional quality (Shigehara, 1994; Temple, 2007a). Previous research explored frequencies of teeth with LEH defects and CO between Jomon, Yayoi, Kofun, and historic Japanese people as well as the relationship of these lesions to Harris lines (Koga, 2003). Similar CO and lower LEH prevalence is observed among Yayoi compared to Jomon samples. Jomon groups were, however, compared as pooled data sets to Yayoi samples (Koga, 2003), despite variable economies between regions (Akazawa, 1981, 1982, 1986, 1999). Three (3) speciﬁc hypotheses associated with the biological impact of agricultural economies in prehistoric Japan are tested by this study. These hypotheses are developed in concert with previous research that suggests little change or improvement in nutritional status following the transition to wet rice agriculture in Southeast Asia (Pietrusewsky and Douglas, 2002a,b; Oxenham, 2004; Domett and Tayles, 2007; Douglas and Pietrusewsky, 2007), and more speciﬁcally, Japan (Koga, American Journal of Physical Anthropology 114 D.H. TEMPLE 2003). First, it is predicted that the introduction of wet rice economies will be associated with a reduction in systemic stress in western Japan. Frequencies of teeth and individuals with LEH defects will signiﬁcantly decline among prehistoric Yayoi compared to western Jomon communities. This decline will be associated with the introduction of a predictable, renewable resource base and exploitation of more nutritionally adequate dietary alternatives. Second, the hypothesis that a similar prevalence of teeth and individuals with LEH defects will be observed between Yayoi agriculturalists and Jomon foragers from eastern Japan is also tested. This prediction expects similar stress patterns between these two groups because both cultures practiced intensive subsistence strategies focused on high yield, reliable food sources, and consumed more nutritious fallback foods. Finally, the hypothesis that CO frequencies remained static following the transition to agriculture in prehistoric Japan is tested. This study predicts stasis in CO prevalence between Jomon and Yayoi period samples as each group thrived in contexts associated with elevated CO prevalence, speciﬁcally sedentary agricultural and maritime environments. Jomon samples from eastern and western Japan are combined to test this third hypothesis because similar rates of CO are observed between the two groups (Suzuki, 1998). BIOCULTURAL CONTEXT Jomon Jomon period (13,000–2300 BP in eastern Japan) cultures were part of a 10,000 year foraging tradition in the Japanese Islands (Imamura, 1996a). Jomon foragers were the descendents of Pleistocene nomads who migrated to Japan around 20,000 BP and subsumed preexisting ‘‘knifeblade’’ cultures (Kobayashi, 2005). These later groups were associated with a ‘‘microblade’’ technology that ﬁrst appeared in Hokkaido, likely in relation to the expansion of cultural networks from eastern Siberia (Imamura, 1996a; Kobayashi, 2005). Microblade Paleolithic industries diversiﬁed from Hokkaido into Tohoku around 14,000 BP, evolved into Mikoshiba tradition, and formed the basis for the earliest spread of Jomon culture (Imamura, 1996a). One set of hypotheses surrounding the earliest migrations to the Japanese Islands suggest that the ancestors of Jomon people migrated from Sundaland (Turner, 1990; Hanihara, 1991). These hypotheses are based on a dental morphological complex (i.e., Sundadonty) observed in Jomon people and shared with the early inhabitants of Sundaland (Turner, 1990, 1992; Matsumura and Hudson, 2005; and others). Multivariate analyses of cranial and dental measurements also ﬁnd similarities between the Paleolithic foragers of Japan, Jomon people, and individuals from Sundaland (Hanihara, 1991; Baba et al., 1998; Hanihara and Ishida, 2005). Other multivariate analyses of cranial and dental traits suggest a Northeast Asian ‘‘point of origin’’ for the Pleistocene ancestors of Jomon foragers (Doi et al., 1997; Dodo et al., 1998; Pietrusewsky, 2005; Seguchi et al., 2007; Hanihara and Ishida, 2009; and others). Cold-derived body size among Jomon foragers and the Pleistocene occupants of the Japanese Islands suggest that this region was initially populated by foragers from a colder environment such as Northeast/Central Asia (Temple et al., 2008). Analysis of classic loci and Y-chromosomes derived from the Ainu suggest a Northeast/Central Asian ‘‘point of origin’’ for the ancestors of Jomon people dating to 20,000 BP (Omoto American Journal of Physical Anthropology and Saitou, 1997; Hammer et al., 2006). More recently, ancient DNA analysis of Jomon period skeletal remains from Hokkaido indicate that the Pleistocene ancestors of Jomon people migrated to the Japanese Islands from Northeast Asia (Adachi et al., 2009). There is some disagreement regarding the use of the term ‘‘forager’’ to categorize the subsistence economies of Jomon people (Imamura, 2006; Underhill and Habu, 2006). It is, however, important to point out that the term ‘‘forager’’ does not describe a static subsistence framework. In fact, many ‘‘foraging’’ societies domesticate considerable numbers of plant species and are characterized by variation in mobility and social complexity (Binford, 1981; Harris, 1989; Smith, 2000). In this sense, Jomon people are encompassed within the foraging subsistence spectrum (Ikawa-Smith, 1986; Tsude, 2001; Matsui and Kanehara, 2006; Underhill and Habu, 2006). Broad consumption of cariogenic cultigens is reported during the Jomon period, despite variation in resource availability (Turner, 1979; Fujita, 1995; Todaka et al., 2003; Temple, 2007a). Spikes in carious tooth frequencies are observed following climatic oscillations around 4300 BP indicating a shift in diet across eastern and western Japan (Fujita, 1995; Temple, 2007a). Dietary changes among prehistoric Jomon people are related to the ubiquitous exploitation of a cariogenic food source such as taro or yams and resulted in elevated caries prevalence among Late/Final compared to Middle Jomon people (Fujita, 1995; Temple, 2007a) with exceptions reported on Hokkaido Island (Oxenham and Matsumura, 2008). Increased consumption of these products did not precipitate agricultural economies per se, as the types of energy expenditure, social organization, and caries prevalence expected in agricultural economies are not observed until the subsequent Yayoi period (Sanui, 1960; Inoue et al., 1986; Imamura, 1996a,b; Oyamada et al., 1996; Tsude, 2001; Todaka et al., 2003; Temple and Larsen, 2007). Yayoi Yayoi period (2500–1700 BP) were the ﬁrst wet rice dependent communities in the Japanese Islands (Imamura, 1996a, b). Cranial and dental size and shape varied between Jomon and Yayoi people in association with environment and gene ﬂow (Brace and Nagai, 1982; Mizoguchi, 1986; Hanihara, 1991; Turner, 1992; Nakahashi, 1993; Pietrusewsky, 2006). Yayoi period agriculturalists were the descendents of people from modern-day Korea or northern China who migrated to Japan and interbred to varying degrees with indigenous Jomon foragers around 2500 BP (Brace and Nagai, 1982; Hanihara, 1991; Nakahashi, 1993; Omoto and Saitou, 1997; Hammer et al., 2006; Pietrusewsky, 2006). Migrants from continental Asia during the Yayoi period introduced wet rice agriculture to the Japanese Islands (Imamura, 1996a,b; Hudson, 1999; Tsude, 2001). The earliest dates for migrant arrival correspond with the earliest dates for wet rice production in Northern Kyushu and Southern Honshu Island, speciﬁcally those obtained from the Doigahama, Itatzuke, and Notame sites (Imamura, 1996b). Tool types and irrigation systems that closely resemble those found at farming sites from southern China and Korea also suggest that wet-rice agriculture was brought to Japan by migrant people (Tsude, 2001). For example, the paddy ﬁeld excavated at the Itatzuke site is remarkably similar to wet rice ﬁelds found in southern China and North Korea between 4000 and 3000 BP (Tsude, 2001). 115 JOMON AND YAYOI STRESS Signiﬁcant energy expended on the care of domesticated plants is recorded during the Yayoi period in the form of large-scale agricultural ecosystems based around wet rice farming (Imamura, 1996b; Hudson, 1999; Tsude, 2001). Precipitous increases in carious tooth frequencies follow this economic shift and bear further testament to the importance of carbohydrate-heavy crops during the Yayoi cultural horizon (Sanui, 1960; Inoue et al., 1986; Oyamada et al., 1996; Todaka et al., 2003; Temple and Larsen, 2007). This increase in caries prevalence is associated with the consumption of a starch-heavy carbohydrate, variation in food preparation techniques (i.e., boiling and processing rice), and increased malocclusion among Yayoi compared to Jomon people (Temple and Larsen, 2007). General variation in cranial morphology between historic Japanese compared to Yayoi people suggest in situ behaviorally-related changes in morphology after the arrival of these migrants to the Japanese Islands (Mizoguchi, 1986). MATERIALS All bioarchaeological data (LEH, CO) were collected by the author (DHT). Human skeletal remains recovered from eight Jomon period sites (Fig. 1; Table 1) represent the foraging component of this study. These sites are dated from the Middle (5000–4000 BP), Late (4000–3300 BP), and Final (3300–2500 BP) Jomon period and include remains from the eastern and western regions of Honshu island. Human skeletal materials recovered from four Yayoi period archaeological sites represent the agricultural component of this study (Fig. 1; Table 1). These sites yielded the largest numbers of skeletal remains for all early agricultural sites in the Japanese Archipelago. Wet rice dependence among these groups is supported by archaeological and isotopic studies (Imamura, 1996a,b; Chisholm and Koike, 1999). All sites were dated using pottery chronology and/or radiocarbon methods. The use of pottery chronology is an accurate method to estimate site occupations in prehistoric Japan because these methods have achieved signiﬁcant precision over many years of description and comparison to absolute dating methods (Aikens, 1995; Imamura, 1996a; Habu, 2004; Tanaka et al., 2005). This study compares Yayoi assemblages recovered from western Japan to Jomon assemblages recovered from eastern Japan. Eastern Yayoi skeletal collections are fragmentary and small in sample size due to highly acidic soil. Those in western Japan were preserved in sand-dunes, while those from eastern locations were interred in soil (Imamura, 1996a). Soil in Japan has a low pH level and is responsible for fragmentation and disintegration of skeletal materials interred outside of shell mounds or sand dunes (Imamura, 1996a). Comparing these two groups could be problematic as these comparisons may not measure a ‘‘direct’’ biological impact of wet rice agriculture in eastern Japan. These comparisons are, however, important because Yayoi samples from western Japan represent the earliest wet rice farmers in the Japanese archipelago. In this sense, comparisons between eastern Jomon communities with the earliest wet rice farmers from western Japan will enhance knowledge of the more immediate differences/ impacts in stress associated with this transition. Fig. 1. Map indicating site locations. Jomon: 1) Nakazuma, 2) Kitamura, 3) Yoshigo, 4) Hobi, 5) Inariyama, 6) Tsukumo, 7) Ota, 8) Yosekura; Yayoi: A) Koura, B) Doigahama, C) Kanenokuma, D) Nagaoka. METHODS Age estimation CO prevalence was compared between age groups. Age was estimated on the basis of long bone epiphyseal fusion (Scheuer and Black, 2000) tooth development/ eruption stages (Smith, 1991), and tooth wear scores obtained from premolar and molar teeth (Scott, 1979). Samples were then divided into two age groups based on these estimations. Age Group 1 includes individuals younger than 15 years, while Age Group 2 includes individuals aged 15 years and older. LEH defects LEH defects were recorded as deﬁciencies in enamel thickness appearing as horizontal grooves or pits on mandibular and maxillary anterior, permanent one of these teeth for both adults and subadults with at least one tooth. LEH defect presence was determined by macroscopic observation aided by the use of a magnifying glass (103), natural ﬂuorescent lighting, and a 100-W Toshiba desk lamp. Identiﬁcation of LEH defects follow Skinner et al. (1995) and Guatelli-Steinberg (2003), where adjacent perikymata were compared to possible LEH defects to prevent confusing normal variation in tooth morphology with disrupted enamel production. The minimum limit for LEH defect identiﬁcation was set at the point where horizontal grooves appeared larger than adjacent perikymata under 103 magniﬁcation. The maximum limit includes LEH defects that were clearly visible as furrows of enamel deﬁciency in the absence of magniﬁcation. Methods for documenting intraobserver error in LEH defect recognition are reported in Temple (2007a,b). These methods determined a level of ‘‘substantial agreement’’ between two rounds of observation. American Journal of Physical Anthropology 116 D.H. TEMPLE TABLE 2. Overall prevalence of anterior, permanent teeth with LEH defects TABLE 3. Percentages of individuals with at least one anterior, permanent tooth expressing an LEH defect Group N Teeth % LEH (Y:) P a Group N individualsa % LEH (Y:) P Yayoi (Y) Eastern Jomon (EJ) Western Jomon (WJ) 965 559 495 30.3 36.8 56.7 – 0.05 0.001 Yayoi (Y) Eastern Jomon (EJ) Western Jomon (WJ) 182 164 122 63.1 48.2 64.8 – 0.05 NSb a Probability values for Yayoi compared to eastern and western Jomon. It is important to note that LEH defects are most objectively identiﬁed by increased perikymata spacing using microscopic measurements (Hillson and Bond, 1997). Since this study relies on macroscopic identiﬁcation, the prevalence of LEH defects reported here represents a minimum estimate. LEH prevalence is ﬁrst reported using individual and overall frequencies. These frequencies were calculated using the following methods: Number of anterior, permanent teeth with at least one LEH defect divided by the total number of observed anterior, permanent teeth (overall prevalence); number of individuals with at least one observable LEH defect on an anterior, permanent tooth divided by number of individuals with at least one anterior, permanent tooth (individual prevalence). LEH defects arising from disrupted physiological homeostasis following severe traumatic injuries may bias interpretations by mimicking hypoplastic lesions normally attributable to stress (Hillson, 1996). LEH defects associated with traumatic injuries are rare in archaeological contexts (Goodman and Rose, 1991). Nevertheless, LEH defects associated with systemic stress are possible to differentiate from those arising as a consequence of physical trauma by chronological matching: LEH defects associated with systemic stress are observable at similar chronological positions on tooth antimeres, whereas trauma-induced LEH defects are generally observed on isolated teeth (Hillson, 1996). Antimere refers to side-speciﬁc identical structures in bilaterally symmetric organisms (e.g., right and left mandibular canines). In an attempt to offset the potential for including trauma-induced LEH defects in the study, LEH defect prevalence on maxillary ﬁrst incisor (MxFI) and mandibular canine (MaC) tooth antimeres were calculated. MxFI and MaC teeth were selected in association with previous observations that suggest these teeth are most susceptible to LEH defect formation (Goodman and Armelagos, 1985). Numbers of mandibular canine (MaC) and maxillary ﬁrst incisor (MxFI) antimeres with at least one observable LEH defect on both teeth divided by the total number of observed MaC and MxFI antimeres, respectively were used to calculate the prevalence of antimeres affected by LEH defects. Cribra orbitalia CO lesions were identiﬁed using a 103 magnifying lens under ﬂorescent lighting and a 100-W desk lamp. CO was identiﬁed as sieve-like lesions on the orbital roof. The presence of marrow hyperplasia through the orbital cortex and/or remodeling on the borders of sievelike lesions helped differentiate this condition from postmortem damage. CO was scored following methods reported by Steckel et al. (2002) and described by Temple (2007b). Prevalence of this condition was compared as the total number of individuals with at least one orbital American Journal of Physical Anthropology a Number of individuals with at least one anterior, permanent tooth. b Does not reﬂect a statistically signiﬁcant result compared to Yayoi samples. roof with evidence of CO divided by the total number of individuals with at least one orbital roof. Statistical methods The G-statistic with continuity correction is a more conservative version of the chi-square test in assessing the independence of nominal data (Sokal and Rohlf, 1995). The G-statistic with continuity correction compares the goodness-of-ﬁt between observed and expected cell frequencies and corrects for greater than expected Type I errors (Sokal and Rohlf, 1995). The G-statistic is applied to data where the observed minus expected cell frequencies are greater than the expected cell frequencies or studies that rely on limited sample sizes—small samples conﬂate the numerator in the chi-square calculation and produce unusually large degrees of difference between data sets (Sokal and Rohlf, 1995). G-tests for independence are employed by previous studies using similar sources of data (Temple, 2007a; Temple and Larsen, 2007). RESULTS LEH defects Overall prevalence of teeth with at least one observable LEH defect is signiﬁcantly greater among eastern (G 5 6.9; P 0.05) and western (G 5 95.6; P 0.001) Jomon foragers compared to Yayoi agriculturalists (Table 2). G-values for the overall prevalence of anterior, permanent teeth with at least one LEH defect are elevated for the western Jomon and Yayoi comparison in relation to a 26% difference in teeth affected by LEH defects. Differences in the frequencies of anterior, permanent teeth affected by LEH defects between eastern Jomon and Yayoi samples are less extensive at 6%. Eastern Jomon foragers have signiﬁcantly fewer individuals with at least one anterior, permanent tooth affected by at least one LEH defect compared to Yayoi agriculturalists (G 5 7.91; P 0.05) (Table 3). No significant differences in individuals with at least one anterior, permanent tooth affected by at least one LEH defect are observed between the western Jomon and Yayoi samples (G 5 0.077). No signiﬁcant differences are observed in MxFI antimeres affected by LEH defects between the eastern Jomon and Yayoi samples (G 5 3.2), while signiﬁcantly greater frequencies of MxFI antimeres affected by LEH defects are observed among western Jomon compared to Yayoi groups (G 5 5.5; P 0.05) (Table 4). No signiﬁcant difference in MaC antimeres affected by LEH defects are observed between the eastern Jomon and Yayoi samples (G 5 0.644). Signiﬁcantly greater frequencies of MaC antimeres are affected by LEH defects among western Jomon compared to Yayoi people (G 5 7.8; P 0.05). 117 JOMON AND YAYOI STRESS TABLE 4. Percentage of antimeric tooth pairs expressing LEH defects on both teeth Group N MxFI antimeresa %LEHb (Y:) P N MaC antimeresc % LEH (Y:) P Yayoi (Y) Eastern Jomon (EJ) Western Jomon (WJ) 77 47 28 35.1 21.2 60.7 – NS 0.05 76 23 39 34.2 56.5 61.5 – NSd 0.01 a Number of maxillary ﬁrst incisor antimeres. Percentage of antimeres expressing LEH defects. c Number of mandibular canine antimeres. d Does not reﬂect a statistically signiﬁcant result compared to Yayoi samples. b Several signiﬁcant differences in the prevalence of teeth affected by LEH defects between Yayoi and western Jomon people support the ﬁrst hypothesis of this study. Few differences in teeth affected by LEH defects between Yayoi and eastern Jomon people conﬁrm the second hypothesis. Cribra orbitalia TABLE 5. CO prevalence by age group and time period Age group 1a Jomon Yayoi Age group 2b Jomon Yayoi a Comparisons of CO prevalence are reported for Age Groups 1 and 2 in Table 5. CO prevalence is not signiﬁcantly different between the Middle to Final Jomon and Yayoi agriculturalists from Age Group 1 (G 5 0.013) or 2 (G 5 0.014). These results are consistent with the third hypothesis of this study that predicted similar frequencies of CO would be observed between Jomon and Yayoi period people. DISCUSSION LEH defect prevalence Several signiﬁcant differences in the prevalence of teeth affected by LEH defects were observed between western Jomon and Yayoi samples. Assuming a decline in stress between the western Jomon and Yayoi samples, these results may be related to variation in resource availability and nutritional quality. LEH defects reﬂect inadequate intake of proteins, vitamins, and iron (Wolbach and Howe, 1933; Mellanby, 1939; Schour et al., 1944). Poor intake of these nutrients is associated with atrophy of the enamel organ (Wolbach and Howe, 1933; Schour et al., 1944) and damage to the Tomes’ process (enamel producing organ) of ameloblasts (Witzel et al., 2006). The enamel organ is a condensation of cells that differentiate into ameloblasts (Ten Cate, 1998). Nutritional deprivation causes enamel organ atrophy through vasoconstriction (Mellanby, 1939; Schour et al., 1944). Poor organizational capacity for the differentiation of ameloblasts and odontoblasts from the enamel organ combined with LEH defect formation follows (Mellanby, 1939; Schour et al., 1944). Tomes’ process damage impairs the capacity for the ameloblast to produce enamel (Witzel et al., 2006) and also results in the production of LEH defects. Similar ﬁndings are reported among children in developing countries. Prevalence of children in Guatemala with teeth affected by LEH defects and overall frequencies of teeth affected by LEH defects were increased among groups that lacked adequate nutritional supplementation (i.e., protein and vitamin fortiﬁcations) (Goodman et al., 1991; May et al., 1993). These ﬁndings suggest that poor consumption of essential nutrients is associated with the formation and greater prevalence of LEH defects. b N individuals % Cribra 24 33 50.0 50.1 209 111 8.6 8.8 Age Group 1 includes individuals less than 15 years. Age Group 2 includes individuals aged 15 years and greater. Teeth and individuals with LEH defects are also observed in great prevalence from a variety of contexts with presumably elevated levels of malnutrition, and especially, regions with resource limitations such as famine-era China and the Protohistoric American Southwest (Zhou and Corruccini, 1998; Stodder et al., 2002). Hunter-gatherer people from Southwest Asia have elevated LEH defect frequencies for teeth and individuals in association with seasonal food scarcity (Lukacs and Pal, 1993). These ﬁndings suggest that nutritionally inadequate food and seasonal scarcity were responsible for the elevated prevalence of LEH defects among western Jomon compared to Yayoi people. It is also possible that increased population density during the Late/Final Jomon period may have added pressure on resource availability in western Japan. Population density declined in eastern and central Japan during the Late to Final Jomon period; western Japan did, however, experience signiﬁcant increases in population density during this same time period (Koyama, 1978). This population expansion appears to be part of a broader response to climatic cooling, where migrations into northern and western Japan combined with dietary change are noted, rather than increased reproductive output within a stressed community (Hudson, 1999; Habu, 2004; Temple, 2007a). Age compositions of deer hunted by Jomon people changed to include large quantities of younger animals during the Late/Final Jomon period suggesting that increased population density was associated with overuse of environmental resources (Koike and Ohtaishi, 1985; Koike, 1992). Similar trends are observed in gastropod exploitation patterns, where range of shell heights expanded to include smaller organisms during the Late/Final Jomon period (Koike, 1986). Finﬁsh consumed by western Jomon people were subjected to seasonal depletion (Kobayashi, 2005), while fallback foods were nutritionally poor (Akazawa, 1986). This suggests that increased population density during the Late/Final Jomon period in western Japan may have encroached on the carrying capacity of an already limited environment and exacerbated nutritional status. American Journal of Physical Anthropology 118 D.H. TEMPLE In contrast, Yayoi people were reliant on a predictable and renewable source of food in the form of wet rice economies (Imamura, 1996a,b) and consumed nutrient rich fallback foods (Chisholm and Koike, 1999; Takahashi, 2009). The earliest form of wet rice produced in prehistoric Japan was resistant to cold weather and able to survive drastic seasonal shifts in temperature (Tsude, 2001). In addition, the form of paddy-ﬁeld agriculture imported to the Japanese Islands was at an advanced stage, having evolved in Northeast Asia over 3,000 years before arriving in this region (Tsude, 2001). Yayoi people also supplemented this diet with maritime resources (Chisholm and Koike, 1999; Takahashi, 2009) and even cultivated various species of freshwater ﬁsh (Nakajima et al., 2009). It is, therefore, likely that the lower prevalence of teeth with LEH defects among Yayoi people is attributable to resource stability and nutritional quality. Similarity in systemic stress prevalence between the Yayoi and eastern Jomon people is associated with exploitation of consistently available subsistence bases and wide dietary breadths. Hypotheses by Yesner (1994) predict that resource abundance and dietary breadth are associated with systemic stress patterns among prehistoric foragers and incipient agriculturalists. For example, patterns of LEH defects in Southeast Asia indicate continuity between foragers and agriculturalists in association with continued broad-spectrum subsistence economies following the transition to wet rice agriculture (Pietrusewsky and Douglas, 2002a,b; Douglas and Pietrusewsky, 2007). Eastern Jomon people relied on a wide dietary breadth with consistently available, calorically dense resources (Akazawa, 1999). Yayoi people continued this trend, incorporating maritime and terrestrial game into agricultural economies (Terasawa and Terasawa, 1981; Chisholm and Koike, 1999; Nakajima et al., 2009). To be certain, Jomon people from eastern Japan and Yayoi groups relied on different foods and resource procurement/production systems. Both groups were, however, active participants in economies characterized by intensive subsistence behavior and the exploitation of calorically dense resources. Cribra orbitalia and parasite infection Similar frequencies of lesions associated with CO are observed between Jomon and Yayoi samples in both age groups. Prevalence of CO was originally thought to form a straightforward relationship with diet, speciﬁcally iron status (El-Najjar et al., 1976). These relationships were later questioned on the basis of various parasites found in the coprolites of prehistoric North Americans (Reinhard, 1990). Cranial porosity also reached signiﬁcant levels among maritime foragers from the California coast (Walker, 1986) and densely populated desert agriculturalists (Walker, 1985) further indicating that dietary factors are only loosely associated with CO. These lesions are, in populous maritime and agricultural environments, associated with iron deﬁciency anemia stemming from parasitic infection, prolonged weaning due to poor resource availability, and diarrhea caused by the use of contaminated water supplies (Walker, 1985, 1986, 2006). A recent survey does, however, suggest that iron deﬁciency anemia alone will not produce the hypertrophic marrow expansion observed in cases of CO and that these lesions are more likely attributable to other nutritional deﬁciencies or infectious conditions of the eye orbit (Walker et al., 2009). Primary diseases contributing to American Journal of Physical Anthropology the development of CO include scurvy, rickets, megaloblastic anemia (MA), and trachoma (Schultz, 2001; Wapler et al., 2004; Walker et al., 2009). It is difﬁcult to differentiate these conditions without histological (Schultz, 2001) or radiographic (Ortner, 2003) sections. It is, however, possible to ‘‘infer’’ speciﬁc conditions associated with CO based on environmental context and lesion distribution (Larsen and Sering, 2000). Rickets is a pathological condition associated with a lack of vitamin D intake (Ortner, 2003). Humans cannot independently synthesize vitamin D and must obtain this nutrient from environmental surroundings. Vitamin D is available in a variety of foods, but in most cases, humans directly synthesize adequate amounts of this nutrient from sun exposure (Ortner, 2003). Under circumstances where sun exposure is negligible, for instance among indigenous foragers of Siberia and Northern North America, consumption of maritime products provide an adequate source of vitamin D (Jablonski and Chaplin, 2000). The climate of Japan is temperate (Honshu Island and regions south) and continental microthermal (Hokkaido) (Fukui, 1977). These regions experience adequate sunlight for vitamin D synthesis outside the context of industrialized or urban environments (Lips, 2007). This suggests rickets arising from dietary/climatic factors was an unlikely contributor to the observed patterning of CO lesions. Rickets is, however, observed in cases of parasite infection (Walker et al., 2009), and therefore, cannot be effectively ruled out as a contributor to these lesions. Scurvy is another diagnostic option for these lesions. Scurvy is a pathological condition attributed to insufﬁcient levels of vitamin C (Aufderheide and RodriguezMartı́n, 1998). Considerable levels of vitamin C are found in citrus fruits, vegetables, and shellﬁsh (Rivers, 1987). Signiﬁcant consumption of shellﬁsh as well as a wide variety of vegetables is reported among Jomon people (Koike, 1986; Crawford, 1992, 2006; Matsui and Kanehara, 2006). Some level of caution is, however, necessary in ruling out scurvy as a diagnostic option. Surprisingly elevated frequencies of scurvy were observed in early historic Florida—a region with abundant access to citrus fruits (Ortner et al., 2001). Underlying infection by parasites and infant diarrheal disease may also signiﬁcantly deplete vitamin C and result in scurvy (Walker et al., 2009). As a consequence, scurvy cannot be effectively rejected as a diagnostic option for the observed lesions. The marrow hyperplasia of CO is also attributable to megaloblastic anemia (MA) (Walker et al., 2009). MA is associated with inhibited DNA production in red blood cells and produced by poor levels of vitamin B-12 or folic acid (Antony, 1995). This condition is rare in adults, but children lacking signiﬁcant stores of vitamin B-12 are particularly susceptible to the disease as are individuals exposed to parasitic infection (Walker et al., 2009). This suggests that MA also contributed to the patterning of CO observed by this study. It is important that parasites speciﬁc to prehistoric Japan be considered as contributing factors to cranial porosity induced by MA, rickets, or scurvy. These conditions include but are not limited to schistosomiasis, trichuriasis, roundworm (Ascaris), and Diphyllobothrium infections. In Asia, schistosomiasis is a disease associated with infection by the cercariae of Schistosoma japonica that survive on snails living in brackish water, depart this vector during daylight hours, penetrate 119 JOMON AND YAYOI STRESS human skin, and disseminate to various organ systems (Mascie-Taylor and Mohamed, 1995). Eggs of Schistosoma japonica are re-released into surrounding environments by urination, mature into cercariae, reattach to snails, and perpetuate new rounds of infection. Environmentally, Schistosoma japonica infections increase in concert with the square acreage of wet rice ﬁelds in rural China suggesting a link between paddy ﬁeld agriculture and schistosomiasis (Yi-Xin and Manderson, 2005). Schistosomiasis inducing parasites are also spread through contact with or consumption of unsanitary water (Mascie-Taylor and Mohamed, 1995). Trichuriasis is a pathological state caused by the consumption of eggs laid by Trichuris trichiura or Trichocephalus trichiuris in dry areas, speciﬁcally those found associated with stored rice and other grains (Stephenson et al., 2000). The disease is also related to accidental ingestion of soil contaminated by the eggs of Trichuris (Stephenson et al., 2000). These eggs hatch in the small intestine, with larvae and mature worms moving into various regions of the digestive system (Crompton and Nesheim, 2002). Subsequent spread through contact with stool is reported (Stephenson et al., 2000; Crompton and Nesheim, 2002). Roundworms are another potential contributor to nutritional depletion and CO distribution among the Jomon and Yayoi people. Parasitic roundworms include members of the genus Ascaris. Ascaris infection occurs when pigs consume the meat of infected rats and humans consume the meat of infected pork, especially in circumstances where meat is raw or undercooked (Aufderheide and Rodriguez-Martin, 1998). Immature larvae are deposited into the small intestine with later migration into the large intestine by mature parasites. Mature parasites then reproduce in the large intestine, depositing larvae into the intestinal mucosa. Finally, infection by the Diphyllobothrium genus of maritime parasite is reported in a variety of coastal contexts (Reinhard, 1990; Bathurst, 2005a,b). In Japan, Diphyllobothrium latum and nihonkaiense infect hosts following consumption of raw or undercooked ﬁsh, with mature organisms migrating to the small intestine (Scholtz et al., 2009). Parasite loads are particularly heavy among maritime foragers, where contact with contaminated, stagnant water and consumption of undercooked ﬁsh provide vectors for infection and reinfection (Reinhard, 1990; Bathurst, 2005a,b; Walker, 2006). Jomon people occupied coastal, populous environments and procured maritime resources on the open water (Koyama, 1978; Imamura, 1996a). Fishing behavior, maritime resource consumption, and exposure to contaminated water supplies likely acted as a primary source of Diphyllobothrium and Schistosoma infection. Jomon people also consumed pig meat (Hongo et al., 2006), and as a consequence, were likely exposed to Ascaris parasites. In addition, Jomon people stored foods including various acorns and walnuts in underground storage units (Imamura, 1996a). Food storage may have placed Jomon people at risk for infection by Trichuris. In fact, whipworms (Trichuris) were observed in coprolites from the Jomon period Sannai Maruyama site (Matsui et al., 2003). The totality of these ﬁndings indicate that CO patterning among Jomon people reﬂect nutrient deﬁciencies brought about by parasite infection, with speciﬁc reference to Trichuris species. Yayoi people stored wet rice as dry grain (Imamura, 1996a,b), continued to rely on maritime resources, and consumed pigs (Aikens and Akazawa, 1992; Chisholm and Koike, 1999). These behaviors likely exposed Yayoi people to Ascaris, Diphyllobothrium, Schistosoma, and Trichuris infection. It is also possible that infection by parasites such as Schistosoma japonica were transmitted through contact with stagnant water in wet rice ﬁelds. More speciﬁcally, however, whipworms (Trichuris) and roundworms (Ascaris) were found in coprolites recovered from the Ikegami-Sone Yayoi site (Matsui et al., 2003). This suggests that the patterning of CO observed among Yayoi people was associated with parasite infection, particularly Trichuris and Ascaris species. It is also important to consider trachoma as a possible contributor to these lesions (Wapler et al., 2004; Walker et al., 2009). Trachoma is an infectious disease of the eye caused by direct contact with the bacteria Chlamydia trachomitis (Wright et al., 2008). Elevated frequencies of trachoma are reported in dense populations lacking access to sanitary bathing and drinking facilities as well as those where young people lack access to clean water for proper facial cleansing and sleep near infected individuals (Wright et al., 2008). Access to sanitary bathing and drinking facilities were a major environmental hazard for prehistoric coastal communities, particularly those with dense populations (Walker, 2006). Both Jomon and Yayoi people occupied regions of high population density (Koyama, 1978). Poor sanitation among these groups is indicated by reports of fecal material within water sources around settlements (Matsui et al., 2003). In this sense, both Jomon and Yayoi people occupied environments where trachoma was a likely environmental hazard and an additional contributor to the CO lesions observed by this study. Stress and the agricultural transition The results presented here differ from previous research addressing patterns of systemic stress during the agricultural transition (i.e., Cohen and Armelagos, 1984; Larsen, 1987, 1995, 1997, 2002, 2006; Cohen and Crane-Kramer, 2007), but are consistent with reports from Southeast Asia (Oxenham, 2000; Domett, 2001; Pietrusewsky and Douglas, 2002a,b; Domett and Tayles, 2007; Douglas and Pietrusewsky, 2007). A general decline/continuity in systemic stress is noted with regard to prevalence of teeth with LEH defects, while CO frequencies remained static. Two major reasons for these systemic stress patterns are important to note. Wet rice agriculture arrived in Japan as a fully functioning subsistence strategy and was supplemented by a sophisticated system of maritime exploitation. The development of agricultural ecosystems implies a fairly long-term process of human/environmental interaction with early systems subjected to failure due to climatic or other environmental ﬂuctuation (Harris, 1989, 1996). The earliest wet rice ﬁelds in Japan best resemble those from China and have signiﬁcantly more complex features than the earliest paddy sites such as Caoxieshan (6000 BP) (Imamura, 1996a,b; Tsude, 2001). In fact, Yayoi paddy ﬁelds post-date the earliest development of wet rice paddies in China by 3,000 years (Takahashi, 2009) and exemplify modiﬁed versions of these earlier sites (Imamura, 1996b; Tsude, 2001). It is, in this sense, likely that Yayoi period agriculturalists did not experience stressors associated with poor control over environmental resources. Instead, these early agricultural people enjoyed the beneﬁt of a well developed subsistence system. It should be noted American Journal of Physical Anthropology 120 D.H. TEMPLE that systemic stress experienced as a consequence of introducing this behavioral strategy into new landscapes may have been experienced, though stress prevalence among Yayoi people still did not exceed levels reported for eastern or western Jomon people. In addition, Yayoi period agriculturalists continued to exploit maritime products following the agricultural transition. Maritime resources provide an excellent source of nutrition, particularly when mixed with cereal products (Layrisse et al., 1968; Larsen, 2003; Murphy and Allen, 2003; Rivera et al., 2003). Prehistoric foragers reliant on terrestrial mammals experienced an increase in the frequency of teeth affected by LEH defects after the transition to intensive plant-based agriculture, in part due to dietary stress (Cook, 1980, 1981, 1984; Lukacs, 1992; Larsen et al., 2002; Hutchinson et al., 2007). In contrast to dietary shifts experienced during the agricultural transition in the New and Old World, there is ample evidence suggesting continued reliance on the procurement of maritime resources among Yayoi people. Similarities in ﬁshing tool kits are noted between Jomon people from eastern Japan and a number of Yayoi period sites (Aikens and Akazawa, 1992; Imamura, 1996a,b; Takahashi, 2009). Stable isotopic analysis of human remains indicates that Yayoi people consumed greater amounts of maritime products than early agricultural people from North America (Chisholm and Koike, 1999). Additionally, previously mentioned studies report evidence that Yayoi people actually cultivated some species of fresh-water ﬁsh (Nakajima et al., 2009). CONCLUSIONS Jomon foragers from western Japan had signiﬁcantly greater frequencies of teeth with LEH defects than Yayoi agriculturalists. Reduction in the prevalence of teeth with LEH defects following the agricultural transition is likely related to the introduction of a predictable, renewable subsistence base into western Japan combined with a system of exploiting more nutritious fallback foods. More speciﬁcally, Jomon foragers from western Japan were reliant on calorically dense ﬁnﬁsh, but consumed nutritionally poor fallback foods. The introduction of wet rice agriculture provided this region with an economic base that was high yield, renewable, cold resistant, and supplemented by maritime foods. Similar frequencies of teeth with LEH defects were observed between Jomon foragers from eastern Japan and Yayoi agriculturalists. The similar prevalence of systemic stress observed between these two groups is attributable to the fact that eastern Jomon and Yayoi people both had signiﬁcant control over environmental resources and relied on nutritionally rich fallback foods that included terrestrial mammals and maritime resources. Finally, CO prevalence between the two groups remained stable. Both Jomon and Yayoi people were exposed to environments that were hospitable to organisms such as Schistosoma japonica, Trichuris, Diphyllobothrium, and roundworms (Ascaris). Trichuris and Ascaris parasites were, in fact, observed in Jomon and Yayoi coprolites. In addition, both groups occupied environments where exposure to bacteria such as Chlamydia trachomitis was likely signiﬁcant. This suggests parasite induced malnutrition and bacterial infection was responsible for the observed pattern of CO among Jomon and Yayoi people. American Journal of Physical Anthropology These results depart from general models of PostPleistocene human evolution that suggest increases in systemic stress followed the agricultural transition. Two primary reasons for the trend in Japan are reported. First, wet rice agriculture was based on a well developed system of paddy farming that evolved over 3,000 years before arriving in the Japanese Islands. Second, Yayoi people continued to rely on maritime products as a dietary source. In contrast, early agricultural people from other regions of the globe relied upon newly developed subsistence systems and nutritionally inadequate cultigens. It is also important to note that Jomon and Yayoi people represent distinct biological groups with unique migratory histories (see references above). Variation in systemic stress between these groups may also represent differences in genetic capacity to buffer against environmental perturbations (see: Hoover and Matsumura, 2008). Research addressing ﬁnite aspects of stress including chronology, duration, and periodicity may further resolve this issue (see: Temple, 2008). ACKNOWLEDGMENTS Masato Nakatsukasa, Kazumichi Katayama, Naomichi Ogihara, and Wataru Yano provided access to and assistance with the Jomon skeletal material housed at Kyoto University. Gen Suwa, Souichiro Mizushima, and Aiko Saso provided access to and assistance with the Jomon skeletal material housed at the University Museum of the University of Tokyo. Takahiro Nakahashi, Yoshiyuki Tanaka, Shozo Iwanaga, Yuji Mizoguchi, and Kenji Okazaki provided access to and assistance with the Yayoi skeletal material housed at Kyushu University. Yoshitaka Miyauchi and Reiko Oomi from Toride Maizobunkazai and Masatsugu Hashimoto from Tokyo Dental College provided the access to and assistance with the Nakazuma Jomon skeletal material. 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