‘Women and Exercise: Physiological Aspects, BARBARA L. DRINKWATER, Php. In 1973 Miki Gorman’s world record time for the women’s marathon was 2:46:36. In 1983 Joan Benoit finished first in the Boston mar- athon in a world record 2:22:42. A good deal has happened in wom- en’s sports during that 10-year period. The enactment of Title IX legislation, combined with the onset of the jogging boom, dramatically changed the role of sport and exercise for females in our society. ‘Title IX ensured that girls and young women would have oppor- tunities to participate in organized sports in our educational system. The acceptance of jogging as a popular pastime provided older women, regardless of their athletic skills, with a vigorous activity for fun and fitness that was socially acceptable. In a sense, the entire country has become a laboratory for testing the popular hypothesis that the differences in male and female responses to exercise ob- served prior to the 1970s could be ascribed in part to a cultural bias that discouraged females from participating in sports, particularly sports that require endurance, strength, and power. In addition to providing new evidence about women’s capacity for maximal per- formance, the active female of today has forced a reconsideration of opinions about her ability to tolerate environmental stressors, her potential for improving fitness with training, and her ability to match her body composition to the demands of her sport. The popularity of the woman athlete as a research subject has produced an extensive list of publications since an earlier review [36] of women’s physiological response to exercise. Space constraints will limit the content of this review to studies published between 1973 and 1983 that deal with postpubertal and adult females. The term athlete will be used to describe any woman who regularly trains for and participates in sport; competition may or may not be involved. Elite athletes, on the other hand, are athletes who are also members of national or international sport teams. Since the majority of studies published in the last decade describing women's response to exercise have concentrated on the female athlete, this review mirrors that emphasis. Descriptive studies reporting maximal aerobic power (Vo, max) of female athletes are limited here to those published since 1976 to maximize the potential for assessing the effects of changes that occurred in women’s sport during the early 1970s. Special at- 2122 | Drinkwater tention will be given to topics not covered in the previous review, such as histochemical studies of skeletal muscle, and to areas where new information has cast doubt on previous assumptions about wom- en's response to exercise. AEROBIC POWER Twenty-five years ago there existed the pervasive idea that“... at puberty development of ability for strenuous exercise stops or even declines in girls while it continues to advance in boys” [85]. A good deal of research effort was directed toward finding the physiological mechanisms underlying the presumed qualitative and quantitative differences between the sexes in endurance capacity [36]. The prob- lem, as we now know, was lack of training rather than an innate biological difference. In the last decade data collected from athletes in many sports demonstrate that women can improve their capacity for “strenous exercise” long past the age of puberty (Table 1). ‘The most important variable in determining the aerobic power (Vo, max) of the woman athlete is the aerobic demand of the sport and pre- sumably, the training program designed to meet that demand. As with males, the highest values for Vo, max are found for distance runners and cross-country skiers. Age is less important—runners in their late 30s have higher levels of Vo, max than swimmers in their teens. On an empirical level, the large number of women of all ages who compete successfully in road races, marathons, and even ultra- marathons is evidence that gender need not dissuade women from participating in endurance sports. Aerobic power represents a blend of biological potential and the physical demands characteristic of one’s life-style, although the rel- ative contribution of each is unknown. In the past it was assumed. that biological differences between the sexes were primarily respon- sible for the observed differences in maximal aerobic power between men and women. It was noted that the difference diminished when Vo, max was expressed relative to body weight or lean body mass (LBM), but in most cases values for men remained above those for women [36]. Since few women had had the same athletic opportun- ities as males, it was suggested that the remaining difference might represent a training effect rather than a sex-linked difference. The events of the past 10 years have provided physiologists with data to test that assumption. Sparling [122] used a meta-analysis of 13 studies to identify the contribution of body size and percentage of body fat (%BF) to the difference in Vo, max between men and women. Expressing Vo, max as L + min-!, ml - kg-!- min-!, and ml - kg LBM~! = min-!Women and Exercise: Physiological Aspects | 23 (68 = = 61s 149 S291 9 Hunasueunoy, [e8t ‘Tot ‘82) estl 98 00g i) OLLE 8b TegAaTO, logt) Lee 6LL ee Leg e'e91 & (86) OTOL 308 O'8F O'Rg L891 OL squua.|, log) eal 861 Lag gg gor I ouidiy {601 ‘801 ‘09 SPaI 861 seg Lg of p4ON, Sunny (ez) 901 ser ca weet 6 Patd uojyenuag (re) $96 L61 res om S 063 BuyaaiwsC (arn) 69IL 68 rls L 08s assouoe] (sn) na a oss 4 10% Buypody (ea) 66 9st #99 st ¥61 qaysed, [sor ‘261 B26 961 gS oe e81 Furuuiss pazuosysuds [a1] 68L ol OLE 3 gle (29 ‘8] Olat 361 o1g 18 Suyuruaag [6a] ves LLL rer ol 6b-0F [6st tal ‘g¢] ror 181 99S ag 68-08 189 “BE ‘06 ‘RB “I tor 681 tes os 62-08 (ra 826 861 os £0s OF OZ-¢1 —- Huruuns aourisiq (s)aaunog (outa) (aru aad steaq) (yum, By. qu) By) (ws) u ay nods Sala ‘xeu ay xew YH xeurt 404, qysiIeM 48H aBy pun pods & saaryry aru fo sasuodsay pondopoiskyg pouaxn yy pun samswaqrany:) [ORKYe UaTaVL24 | Drinkwater resulted in gender differences of 56, 28, and 15 percent, respectively. For men and women training for the same sport at equivalent levels of competition and tested in the same laboratory, the differences between the sexes averaged 51.5, 18.6, and 9 percent (Table 2). Con- trary to earlier reports (4, 36], Sparling [122] found that differences between trained men and women (25%) were less than between un- trained groups (30%) when Vo, max was calculated relative to body weight. The estimate of 18.6 percent difference for the athletes listed in Table 2 also represents an improvement over the 20-25 percent difference reported in 1973 [36]. However, any substantial difference will put women at a disadvantage in weight-bearing activ- ities. The caloric cost of running, for example, is the same for men and women when energy expenditure is related to body weight [30, 47, 58], The runner, male or female, with the higher Vo, max TABLE 2 Effect of Body Size and Composition on the Magnitude of Differences in Maximal Aerobic Power (VO, max) Between Male and Female Athletes Vo, max. : Vo, max (nl « kg Voz max (ml: kg’! - Mol Sport, Sources Age %BF (L-min) — min-!) min?) Males Skiing Alpine 218 © 10.2 5.03 743 Nordic 22.7 79 5.34 78.9 Nordic 25.6 10.2 542 89.8 Distance running 264 114 431 69.8 nis 420 163 3.87 60.1 Volleyball 26.1 12.0 4.80 63.8 Orienteering 312 163 4.45 73.6 Females Skiing Alpine 19.5 20.6 3.10 52.7 66.5 Nordic 20.2 © 15.7 3.44 154 Nordic 243° 218 4.03 86.4 Distance running 25.8 18.9 2.82 68.9 Tennis 39.0 20.3 354 Volleyball 216 © 179 3.57 61.6 Orienteering 29.0 18.7 2.68 36.7 Percent Difference ((M — F)F] x 100 Skiing Alpine (60) 62 26 12 Nordic [60] 55 19 5 Nordic [108] 34 15 4 Distance running (28] 33 u 1 Tennis (130] 58 4 8 Volleyball [101] 34 uM 3 Orienteering [74] 66 34 30 GBF = percent body fat.Women and Exercise: Physiological Aspects. | 25 (ml - kg! « min-!) will have the advantage of running ata lower %VO, max at any speed. While some women do have higher levels of Vo, max than some men, among elite endurance athletes the male usually has the advantage. The highest level of aerobic power reported for a woman athlete is 77 7 O, « kg! + min=!; for a male the record is 94 ml O, - kg-! - min-!, a difference of 22 percent [10]. When the contribution of body fat to sex differences in Vo, max is removed mathematically, the remaining difference must represent either an actual difference in physical conditioning or a sex-linked difference in the capacity to transport and utilize oxygen. There are practical as well as theoretical reasons for identifying the extent of biologically determined differences in the maximal aerobic power of men and women. Since the functional capacity of the oxygen trans- port system can be a confounding factor in physiological research, it is important that the investigator be able to control this factor by equating subjects on fitness levels. Cureton [27] discusses this problem at length and recommends matching men and women on activity histories and Vo, max (ml - kg LBM~! - min-!). This technique and the residualized Vo, max scores recommended by Katch [71] are two practical methods of assuring that varying levels of cardiorespiratory fitness do not bias experimental data. However, an exact match on Vo, max (ml - kg LBM~! - min-!) would ignore the probability that other sex-related variables affect Vo, max. Cureton [27] suggests a difference of about five percent in Vo, max (ml - kg LBM-! - min=!) can be attributed to inherent biological differences between the sexes. If the difference between male and female subjects is between five and ten percent, one could be reasonably confident that the groups are matched for physical fitness. Further investigation will be nec- essary to determine how much of the remaining difference can be attributed to the lower hemoglobin concentration of the female, her smaller muscle mass—to—body weight ratio, or the smaller compo- nents in her oxygen transport system. Even though the average Vo, max values for female athletes (Table 1) have not increased dramatically since 1973 [36], some individuals and some teams have values well above the average. Women runners and Nordic skiers have attained scores over 70 ml - kg~! - min-' (10, 139], and the average for five members of the Finnish Nordic Ski Team was 68.2 ml: kg! - min-! [108]. There has also been a 30 percent improvement in aerobic power of women basketball players, no doubt representing the increased aerobic demands of the game now that women are playing the full court (127]. Perhaps the most heartening aspect of the change in attitude to- ward women’s participation in sport has been the number of women who choose to remain active as adults. In 1973 there were no data26 | Drinkwater on women athletes beyond the age of 25 [36]. Now it is possible to compare older female athletes (Table 1) with their less active or se- dentary counterparts (Table 3). Women in their 30s and 40s who run [58, 126, 139], play tennis [130], or climb mountains [39] have higher levels of cardiorespiratory fitness than nonathletic women in their 20s. For athletic older women a marked aging effect on aerobic power is postponed until the fifth decade. Since there are only 10 women in the 40- to 49-year-old group in Table 1, the sudden dec- rement in Vo, max at that point may be a sampling artifact. More data on masters athletes will provide a clearer picture of the extent to which vigorous exercise minimizes the aging effect on the endur- ance capacity of women. : Among a general population of females a decrement in Vo, max begins in the 20s (Table 3). Several cross-sectional [38, 65, 100] and longitudinal [2, 5, 96] studies have attempted to define the rate of that decline by regression analysis of data from both active and se- dentary women. Asmussen et al. [2] and Astrand et al. [5] retested former physical education students after 40 and 21 years, respec- tively. Regression slopes across 40 years were calculated as — 0.63 ml O, + kg! min-! - yr-! [2], and for 21 years as —0.44 ml O, - kg-! min-! - yr-! [5]. The original levels of aerobic power for the women in both studies were well above average. Therefore, the decline in Vo, max with time probably included detraining as well as aging effects. In contrast, Profant et al. [100] reported minimal change with age from the fourth to the seventh decade—a decrease of only 0.2 ml - kg-! + min-!- yr-!. Their younger subjects averaged 28.91 ml O, « kg-! = min-! in Vo, max, which probably explains the minimal change across age. Differences in apparent aging effects between longitudinal and cross-sectional studies raise questions about the validity of both meth- ods. Plowman et al. [96] were able to study longitudinally a group of women who had previously been part of a cross-sectional study of age and aerobic power [38]. Slopes for four of five age groups did not differ significantly from the original regression line of Vo, max versus age. The exception was the youngest group, ages 20 to 30 in the original study, which showed no change in Vo, max over a 4.6- year period. The longitudinal changes in maximal heart rate (HR max) also corresponded to the best-fitting curve for HR max versus age in the original study. The authors concluded that cross-sectional studies can provide valid information about the effect of aging on response to maximal exercise. The combined slope for all age groups in the longitudinal study [96] was — 0.323. For the age groups 20 to 60 in the companion study [38], the slope was — 0.291. Both figures27 Women and Exercise: Physiological Aspects rey Apoq uasiad = 40% {11 ‘¢9 “gel 99g S9sI 6s = eto Geer. ee 49a0 pur Og (oor ‘s9 ‘88 gag 6LLI oe e418 «689 SPOT aS Aaewapas loot ‘8s] £09 VLet 18% es eee) Opal ca aatloy 69-05 (oot ‘s9 ‘8¢) ag BOLI £98 Glo eed 0991 ra Aaeuapag. loot ‘gs] 988 zee! a8 Loe «09 HOT’ aanoy 6r-0F {981 ‘OOL ‘29 ‘8s] FOL Vest L6% Ole yo bel 211 Aaeruapas, (oot ‘s¢] Ves oret PLE 1s ag LUO at any 68-08 (F21 ‘F6 ‘08 i “LL ‘99 ‘bF ‘88) eee s¥61 one FIS «BRS MRT BIT Aaeiuapas (az ‘86 “FE ‘BU) 88 ¥'68I ver 008 SLs sol 98 aay 63-08 ‘s991n0g (cum) (uur aod swag) Guta. , 49% YD) (up) « aay Sd.La “xeU 34 xeut YH xeu 4 1BIEM 1818 uauoy Cenjuapag pun sajazyinuony aanry fo asizsaxq jourxvypy 07 asuodsay pun srysriajrvavycy yorskyd aTaVL,28 | Drinkwater are very similar to the —0.307 slope calculated from 29 studies of women in eight countries [36]. Provided that training state remains consistent across time, it ap- pears that women who are not endurance athletes may expect a decrease of about 0.3 ml O, - kg-! + min! - yr~! as they age. However, it should be noted that linear regression equations may not fully describe some of the changes in physiological functions that occur with age. Some variables, including Vo, max relative to body weight and lean body mass, demonstrate a cubic trend with age—a minimal change for some years and then a rapid decrement [38]. More en- couraging is the possibility of changing a negative slope to a positive one, as many women are doing by embarking on personal training programs. AEROBIC TRAINING ‘There is now ample evidence in the literature to indicate that women of all ages benefit from physical conditioning programs [42, 72, 80, 94, 103]. The changes that occur are similar to those observed for males: an increase in maximal aerobic power and ventilatory volume [44, 80, 94] an increase in oxygen pulse [13, 49, 72, 94], and an improvement in submaximal work performance (31, 42, 49, 59, 75, 80]. Most investigators (80%) report no change in maximal heart rate, although occasionally a significant increase [75] or decrease [26, 31] is noted. The training programs studied varied widely. Frequency ranged from two to six days per week, with occasional two-per-day training schedules for athletes [75, 142]. Intensity was expressed as a per- centage of maximal heart rate, heart rate reserve, or maximal aero- bic power, or was not quantified. The shortest training period was four weeks, and the longest, eight months. The average improve- ment in maximal aerobic power for this assortment of programs was 18.7 percent. The variability in the training effect on Vo, max has been linked to a number of factors, but primarily to the initial fitness level of the subjects. Knowlton et al. [75], for example, re- ported a correlation of —0.75 for initial Vo, max versus percent improvement. The range for the studies cited here was 7.2 to 37 percent. For the studies that reported 13 percent improvement or less, the mean initial Vo, max was 42.0 ml - kg~! - min='; for changes greater than that; the original value was 33.2 ml - kg-! - min-!. When men and women trained together the training response was almost identical [13, 31, 41]. Eddy et al. [41] recruited men and women as subjects for a study of continuous versus interval training, with daily work output constant between methods. Men increasedWomen and Exercise: Physiological Aspects | 29 Vo, max percent, women, 14.2 percent. Other training re- sponses—submaximal and maximal heart rate and lactic acid con- centration—did not differ between sexes or style of training. Burke [13] found two differences between men and women as a result of an eight-week running program. Women lost more weight, and men had larger increases in Ve, max. Changes in Vo, max, oxygen pulse, and heart rate were the same. The arrival of women at West Point provided Daniels et al. [31] with an opportunity to compare the effects of a field training program on men and women who were well motivated and performing under considerable pressure. The women improved Vo, max by 7.9 per- cent, the men, by 2 percent. Both sexes increased Ve max, decreased HR max, and showed a similar improvement during submaximal work. The lack of a significant increase in Vo, max by men was attributed to their high initial level of fitness (59.4 ml - kg~! - min-'). Although the overt response to training is the same for men and women, there has been a suggestion that the mechanism underlying the increase in Vo, max differs between the sexes. Men increase Vo, max by increasing both stroke volume (SV) and Cao, — Cvo, [107]. In contrast Kilbom and Astrand [73] found that women improved Vo, max by increasing SV with no change in Cao, — C¥o,. More recent evidence provided by Cunningham and colleagues [25, 26] suggests that the apparent ference in response was not due to gender but to the intensity of the training stimulus. When young women were trained at a high intensity (70 and 100% Vo, max) for 12 weeks they too increased Cao, — C¥o,. The authors [26] conclude that the stimulus for peripheral adaptation is a high-intensity exercise program, and that men and women respond equally to that stimulus. MUSCLE FIBER The needle biopsy technique has allowed investigators to use histo- chemical and biochemical methods to examine muscle fibers from men and women and to identify similarities and differences in fiber composition and metabolic potential. As data have accumulated from different sports, efforts have also been made to determine how fiber type or enzyme activity differs between endurance and nonendur- ance athletes. Most studies report fiber composition by designating the percentage of fibers adapted for prolonged aerobic work, the slow-twitch (ST) group. The remaining fibers, those designed pri- marily for strong rapid contractions, are identified as fast-twitch (FT) and sometimes further subdivided into fast-twitch oxidative-glycolytic (FOG) and fast-twitch glycolytic (FG). Since the demarcation between30 | Drinkwater FOG and FG is not precise in humans, most investigators simply report percentage of FT [23]. Differences in fiber composition are more marked between en- durance and nonendurance athletes than between the sexes [23, 99, 110]. The percentages of ST fibers for untrained men and women are 52 percent (15, 23, 108] and 51.2 percent (8, 19, 26, 99, 110}, respectively, while the mean for women athletes varies from 27.4 to 60.6 percent (Table 4). With women as with men, it is the endurance athlete who has the largest percentage of ST fibers [23, 24]. Where data are available for men and women athletes in the same sport, there is no significant difference in percentage of ST fibers between sexes (99, 110]. The gender difference appears in the size of the fibers (Table 4). Among untrained individuals, women have 68 and 71 percent of the male FT and ST fiber areas, respectively, a figure closely ap- proximating the sex difference in strength. Although endurance and nonendurance male athletes have larger fiber areas than female ath- letes in similar sports, the percentage differences vary according to the demands of the activity. In events relying primarily on anaerobic energy and strength, women athletes have 66 and 71 percent of the male FT and ST fiber areas, a difference similar to that observed in the untrained group. Women trained for endurance events have 85 percent of the corresponding male areas for both ST and FT fibers. Their ST/FT area, 1.06, is the same as that of male endurance ath- letes. While percentage of fiber types is believed to be determined bio- logically, fiber area can be increased by training [23]. Male sprinters, throwers, and jumpers, for instance, have larger FT areas and smaller ST areas than male distance runners and competitive cyclists [15, 23, 101, 108}. Female endurance athletes exceed women trained in no- nendurance sports in both ST and FT fiber areas. While this obser- vation may simply reflect chance differences due to small sample size, it would be interesting to determine whether there is a sex difference in the effect of high-intensity anaerobic and/or strength training on muscle fiber hypertrophy of the FT fibers. The oxidative potential of muscle fibers is usually represented by a measure of succinate dehydrogenase (SDH) activity. Activity levels are increased by training, and are higher in endurance athletes than in nonendurance athletes of both sexes [15, 108, 110]. Rusko et al. [108] reported equal SDH activity for elite male and female cross- country skiers, but Burke et al. [15] found male cyclists had 50 percent greater SDH activity than female cyclists (Table 4). The women, how- ever, had twice the SDH level of untrained men. A study by Costill et al. [23] of athletes in six track and field events found the direction31 ‘xercise: Physiological Aspects | Women and E aseuafoupayap areisey = Ha'T ‘asedaoydsoyd = Hd !aseuadoapdtap areupons = TCS. SMWIUDOASeH = + ‘yA sMIsKA = “TAL FOL x9 860 S6lb — SL8E 88h DOTS Ol Bes [g3] paurnu, 68 _ oe a a a TAGS G1 OLT [601] Buys Arunes-ss0 zs 69 980 Ices 261g 6:9F Oslg & aKa [ga] snosip pur ioys ool 86 06 90 69cF —-b9RF ag OO € £08 [ga] apasel arol SL rR 880 SI¢ £9IF Orb OLR & ER (gz) duanf y8iy pur dum! 8007 brL = BLOT sol z¥9g 6909 ¥09 9909 £ — 66L [kB] SuMUNs aourrsip-arppyy osel o0B FOL 660 086g = ZeLE 89% Ob % G6r 42] sturdy, 9498 ot $60 gtze Leng Olb TASOS £L0R = fe oa ae'0 $60r —_S0RF = TABRP GRR Hat Hd Has (AALS) dai is (ware %) (%) Staqly ue aay aaanog ‘Lodg, “ar. Ba eV Tumeyoqy aHIs 1s uamogy paurnsjuy pun pourvey fo Grayry aukeug aasnpy pun sous rDMY,) Laqiy Asn baTaVL32 | Drinkwater of differences in SDH activity varied with the event. Women had higher levels than men in the throwing events, but since the vastus lateralis was the sampling site the training effect on SDH activity in the arm and shoulder muscles involved in throwing was not mea- sured. The glycolytic enzymes lactate dehydrogenase (LDH) and phosphorylase (PH) showed no consistent differences between men and women track and field athletes [23]. Women cyclists, however, had higher LDH and PH levels than their male counterparts [15]. Although there is some conflicting evidence on enzyme activity, investigators in this area agree that men and women have similar muscle fiber composition, and that the main differences between the sexes relate to fiber area. In the late 1970s, when much of this work was published, some individuals suggested fiber typing as a means of identifying young athletes with a genetic potential for certain sports. This idea did not stand the test of time. These histochemical and biochemical techniques are research tools for identifying and quan- tifying the function and metabolic potential of muscle fibers. There are still very little data available for women athletes, certainly not enough to profile the successful female athlete as a model for a selection process. ENVIRONMENTAL STRESS The observation by I. Astrand [3] in 1960 that core temperature during exercise was more clearly related to relative (%Vo, max) than to absolute exercise intensity had important implications for experi- mental design in thermoregulatory studies. Unfortunately, her re- port went largely unnoticed until confirmation of her finding was reported by Saltin and Hermansen in 1966 [111]. By then a number of papers reporting gender differences in thermoregulation had ap- peared. In almost every instance women exhibited more evidence of heat strain than men when both performed the same absolute work. Unfortunately, some of these same studies are still being cited as evidence that women cannot tolerate heat stress as well as men when, in fact, they were working at a higher %Vo, max. There are occasions when this type of comparison is justified, as in industrial situations where workers perform the same task regardless of sex. However, when physiological mechanisms are of interest, all confounding vari- ables, such as fitness level, must be eliminated either statistically or by an appropriate experimental design. Assigning the same relative exercise intensity to all subjects does not compensate completely for differences in aerobic power. When athletic women were compared with nonathletic women during ex- ercise at 30% Vo, max in three thermal environments, the indicatorsWomen and Exercise: Physiological Aspects. | 33 of heat strain—core temperature (T,) and heart rate (HR)—were similar in mild heat (28°C, 12.6 torr vp) and hot-humid (35°C, 28.0 torr vp) conditions for both groups [37]. However, in a hot—dry environment (48°C, 8.7 torr vp) the nonathletes were unable to main- tain stroke volume (SV) or cardiac output (Q). The athletes not only maintained Q but were able to increase forearm blood flow markedly over that observed in the less stressful conditions. While T, was the same for both groups, tolerance time was significantly reduced for the nonathletes. ‘To clarify the role cardiovascular fitness plays in temperature reg- ulation, the protocol was repeated using women marathon runners and age-matched controls with similar body surface areas [40]. En- durance training was associated with a lower resting T,, a larger plasma volume, a greater influx of plasma protein into the vascular space during exercise, an earlier onset of sweating relative to time and T,, and a smaller decrease in plasma volume during the exper- iment. The runners also had lower heart rates, T,’s, and mean skin temperatures (T,) throughout the exercise. Kobayashi et al. [76] avoided the problems of the interaction of exercise and heat stress by observing Japanese female athletes and female and male nonath- letes at rest in a 32°C environment with their legs submerged in a water bath at 42°C. The female athletes had lower T,’s and heart rates than the other groups throughout the two-hour period. The T, was the same for all subjects. ‘An obvious problem with these types of studies is the possibility that some factor of self-selection accounts for the ability of these women to endure both strenuous training regimens and heat stress. The hypothesis that physical training might partially acclimatize ath- letes to heat stress has been a subject of controversy for years. Two recent reviews [45, 56] on the topic discuss how variations in exper- imental protocol have led to conflicting results, and conclude that training at more than 50% Vo, max for 8 to 12 weeks in a cool environment does indeed improve heat tolerance for both men and women. Cohen and Gisolfi {21} have gone one step further and shown that interval training at 90 to 95 percent HR max in 22°C not only improves women’s heat tolerance but also decreases the time required for acclimation to exercise in a hot environment (45°C). In essence, the effects of physical training mimic the classic indications of heat acclimation—a decrease in T, and HR, an increase in tolerance time, and a favorable change in the sweating response. Although concern had been expressed that conditioning provided protection for only a limited period of time, two hours or less, Gisolfi etal. [57] reported that male distance runners successfully completed four hours of exercise at 25 to 30% Vo, max in both hot-wet and34 | Drinkwater hot—dry environments. More recently, Cohen and Gisolfi [21] found that after only 11 weeks of interval training four of six women could complete four hours of work at 30-35% Vo, max in dry heat (45°C). However, a word of caution is in order. Women athletes should not assume that they are immune to heat illness or that they are totally heat acclimatized. Most heat tolerance tests use a light exercise in- tensity, less than 50% Vo, max, while most distance runners train and race at a pace exceeding 60% Vo, max. At a high exercise in- tensity, metabolic heat production combined with a high environ- mental thermal load can exceed the athlete’s capacity to dissipate heat, no matter how well trained she is. It is also generally assumed that only athletes whose training regimens result in marked elevations of T. are likely to increase their exercise—heat tolerance [91]. A high Vo, max per se does not guarantee immunity to heat stress. One unresolved question in the training-induced acclimation con- troversy is the response of the sweating mechanism. The group at the John B. Pierce Foundation reports that training modifies the peripheral response, increasing the gain of the sweat rate—versus— core temperature relationship (m,./T,) with no change in the zero point central drive for sweating, while heat acclimation has the op- posite effect [86]. Other investigators [45, 61] hold the opposite view: that exercise training affects the central drive and heat acclimation modifies the peripheral response. The dispute is of particular interest to women because of the persistent belief that women’s low sweat rate relative to men’s places them more at risk from heat. Until this issue is resolved, it is reassuring to note that the effect of training on sweating is similar for men and women: a higher sweat rate at any T, or A°C and an onset of sweating at a lower T, [21]. Investigators agree that when women participate in standard heat acclimation protocols their response is qualitatively the same as that of men [89]. However, according to Burse [16] women are less likely to acclimate successfully, and even if they do they will exhibit greater cardiovascular strain and will sweat less than men. These conclusions are based on a number of older studies that did not match the sexes for cardiovascular fitness or activity levels. More recent studies [6, 63] do not support the earlier contentions. Avellini et al. [6] matched four men and four women on Vo, max, expressed as ml O, » kg LBM~! - min-!, on body surface area (Ap), and on surface area-to-weight ratio (Ap/wt) before acclimating them for 10 days to a hot—humid environment (36°C/30°C, db/wb). In the preacclimation heat tolerance test the males had significantly higher heart rates and core temperature in spite of having a higher overall sweat rate. One woman completed the entire three hour test; the longest period for a man was 170 minutes. After acclimation theWomen and Exercise: Physiological Aspects | 35 response of the two groups was the same for the first 90 minutes. Thereafter T, and HR continued to increase for men but stabilized for women. Again the sweat rate of the men was higher, and the increase in rate with acclimation was greater for the women. The onset of sweating was earlier for the men before, but not after, ac- climation. The authors concluded that women may have an advantage in humid heat, since they can perform the same work as men with less fluid loss and less physiological strain. Frye and Kamon [52] raised the possibility that men’s higher sweat rate might provide them with an advantage in dry heat, where the potential for evaporative cooling would be greater. ‘Their subjects were approximately equal in aerobic power, 56.33 and 54.08 ml O, + kg-! - min-! for men and women, respectively, and in Aj/wt. Acclimation was carried out in a 48°/25°C, db/wb environment for 8 to 9 days by walking on a treadmill for two hours at an energy expen- diture equivalent to approximately 27% Vo, max. Prior to acclimation the men had longer tolerance times and higher m,,. Although final T., Ty, and HR did not differ between the sexes, the increase of T, with time was steeper for the women after 25 minutes in the chamber. After acclimation there were no differences in thermoregulatory re- sponse between the sexes. An interesting observation was the simi- larity in sweating threshold and sensitivity (Am,,./A°T,) for men and women both before and after acclimation. Women increased m,. by 41 percent through acclimation, and men increased rh, 10 percent. As a result postacclimation m,, was the same for men and women. Apparently the hot-dry environment did not provide men with an advantage in heat dissipation once the women had become accli- mated, Horstman and Christensen [63] tested essentially the same hy- pothesis in a similar environment (45°/23°C, db/wb) but at a higher exercise intensity (40% Vo, max). The women evidenced slightly more heat strain during the preacclimation test, although the rates of increase in T, and HR, m,,/AT,, and performance time were the same for both groups. By the sixth day of acclimation the women had significantly longer tolerance times and smaller increases in T.. By this time absolute values for AHR and HR were the same, while the increase in T, was higher for the men. Throughout acclimation m,, was higher for men, but by day 6 and at day 11 m,/AT, was higher for the women. In this study the women acclimated more quickly and completely than the men. Another paper from the Natick group reported opposite results [116]. In a more extreme environment (49°C, 20% rh) and during a shorter acclimation period (6 days) men and women walked on a horizontal treadmill at 1.34 m - s~! for 120 minutes. Since the women.36 | Drinkwater had higher T, and T, before and after acclimation, the authors concluded that women have a higher thermoregulatory set point. The authors acknowledged that the women (Vo, max 40.5 ml - kg~! min-!) were less fit than the men (Vo, max 52.3 ml - kg-! - min-}), but did not consider the difference a major factor in their results. It is interesting that a companion paper [7] reported no difference in T, between the same men and women when the exposure was ex- tended to four hours. A higher HR for the women during this ex- periment was attributed to their higher relative exercise intensity. Although Wyndham et al. [146] long ago suggested that men’s “prolific” sweating was wasteful and that women might have a more efficient sweating response, Frye and Kamon [53] were the first in- vestigators to examine this hypothesis in detail. Four men and four women, equated on Vo, max, were acclimated to humid and dry heat before heat tolerance tests in both environments. Measurements in- cluded total body ,., chest m,., sweat gland activity/Ap, sweat gland flow, and maximum sweat gland activity/Ap, using methocholine. Ef- ficiency of sweating was calculated as a ratio of required to observed sweating. As expected, the men had higher sweat rates than the women in the humid heat, but the women were more efficient, 0.99, than the men, 0.81. There were no gender differences in sweat rate or efficiency in dry heat. The women’s advantage in humid heat was related to their higher Ty, which decreased heat gain from the en- vironment and increased evaporative potential by increasing (Py — P,). The women reduced their sweat production by reducing the number of secreting sweat glands, while the men decreased flow. Since the men recruited only 75 percent of their available glands, compared to the women’s 90 percent recruitment, the authors sug- gested that males may have greater reserve capacity. Although men and women may handle their sweating responses in slightly different ways, the outcome in HR and T, was the same. The preponderance of evidence indicates that the response of in- dividuals to exercise during acute exposure to heat stress depends more on the state of their cardiovascular system than their gender [35, 37, 92, 133]. Following acclimation, tolerance to heat stress is improved. The process is similar for men and women, and usually tends to diminish any preacclimation differences between the sexes. When one group acclimates less successfully than the other [51], the cause may lie in initial fitness levels, since it has been demonstrated that prior conditioning affects the rate of acclimation [21]. Even when resting conditions are used to minimize the effect of physical fitness (11), there are still marked differences between active and sedentary individuals. All of these observations emphasize the importance of selecWomen and Exercise: Physiological Aspects. | 37 the appropriate technique for matching subjects on cardiovascular fitness. Both aerobic power relative to body weight and lean body mass have been used to match men and women (6, 52]. However, if lower hemoglobin levels account for a large proportion of the dif- ference in Vo, max between the sexes, does accounting for differ- ences in body composition alone provide an equable match? Are a man and a woman with Vo, max of 65 ml - kg LBM~! - min equally trained and equally fit? Another area requiring further investigation is the role of mor- phological factors in heat tolerance. In general, women have larger A,/wt than men, and several authors [16, 48, 51] have suggested that this may be a disadvantage in some environmental conditions. Sha- piro et al. [114] reported originally that a higher A,/wt for women might explain their lower tolerance for hot—dry conditions than men. A later paper [115] examined the same data expressed relative to Ap rather than body weight, concentrating on thermal balance and heat transfer. The authors [115] concluded that a higher A,/wt did not place women more at risk in a hot—dry environment, because their higher T,, protected them against excessive heat gains. Some degree of standardization in expressing thermoregulatory responses might resolve some of the conflicts in the literature over gender differences in thermoregulation. Although the evidence is strong that the menstrual cycle has little [6, 11, 112] or no effect [48, 51, 52, 54, 64, 134, 135) on the work- heat tolerance of women, there are some intriguing areas for inves- tigation. The primary indicators of heat strain, HR, T, and m,., are not markedly affected by cyclic hormonal changes (48, 51, 52, 64, 134, 135], but there may be some subtle changes in fluid dynamics [55, 112]. Gaebelein and Senay [55] reported more rapid decreases in plasma volume (PV) during the follicular phase, with attendant rapid increases in hemoglobin and osmoconcentration. The changes were magnified with hypohydration. In spite of the decrease in PV there were no cycle effects on T, or HR. It would be interesting to observe the effect of a heavier work load (> 30% Vo, max) and a longer period of heat exposure (> 60 min) to determine whether differences in T, and HR could be elicited. One of the main problems with this study [55] and others [6, 11, 52, 54, 64] is their small sample sizes. It is understandable that long- term studies requiring repeated heat exposures will suffer from sub- ject attrition, but small n’s diminish the power of the statistical tests. Fortunately, the replication of results from many studies [48, 51, 54, 64, 134, 135] adds weight to the conclusion that monthly fluctuations of hormone levels do not seriously affect women's thermoregulatory response to exercise in the heat.38 | Drinkwater Reports from other areas of investigation related to heat stress also failed to find differences in response related to gender. Men and women doing intermittent work in the heat and resting in a cool room both respond to the weighted mean of the environmental con- dition and metabolic heat production [37]. When hypohydrated prior to heat exposure, both demonstrate similar changes in response [18]. Even the evaporative coefficient is the same for men and women [69, 70). Although Senay and his collaborators [51, 112, 113] have de- tected differences in fluid dynamics between men and women during heat exposure, they face the challenge of showing that these different patterns affect heat strain in men and women who have equal aerobic capacity or who are equally acclimated. BODY COMPOSITION In an earlier volume in this series, Oscai [90] reviewed the literature describing the effect of exercise on weight control. Since then two other critical reviews [46, 138] of research on this topic have been published, so only a brief overview will be given here before dis- cussing the implications of this research for female athletes and for women in general. Available data support the hypothesis that exercise alone or in combination with dietary restriction results in a decrease in body fat. The decrease, however, does not always match the decrement ex- pected from the calculated caloric deficit [46, 138]. The difficulty in measuring caloric intake and energy expenditure precisely is cited as a probable cause of this discrepancy. Even though the percent change in body fat is small, the benefits of adding physical activity toa weight control program may exceed the quantitative results. One of the major problems in diet-induced weight loss is the inability of so many people to maintain the desired weight once they have achieved it. It is encouraging, therefore, to find some studies that suggest that ability to maintain the target weight is enhanced if weight was lost through an exercise-plus-diet regimen [46]. Equally intriguing is the current interest in the role of exercise in appetite control [138]. Some investigators [90] are suggesting that exercise may suppress the appetite through yet unidentified mech- anisms, a reversal of the common belief that exercise increases food intake. Both these hypotheses must be tested extensively before these claims can be verified. However, there is evidence that diet alone results in a loss of lean tissue as well as fat, while exercise alone or exercise and diet plans cause preferential loss of body fat and spare lean tissue [1, 81, 138, 146]. This in itself is sufficient cause to rec-Women and Exercise: Physiological Aspects. | 39 ommend exercise as a tool to any woman concerned with maintaining body fat within reasonable bounds. The definition of “reasonable” depends on the motive of the in- dividual involved. The athlete manipulates her fat content to improve performance and is interested in knowing the optimal percentage of body fat (%BF) for her sport. Distance runners are seldom happy with a fat content greater than 12 percent, while distance swimmers are content with 25 to 30 percent, for buoyancy and as protection against hypothermia. The wide range of %BF shown in Table 5 illustrates the diversity among athletes who represent different sports. In activities where body weight represents the workload, %BF is far below the 25 percent average reported for college women. Even TABLE 5 Percent Body Fat (%BF) for Female Athletes by Sport and Age Sport ‘Age n _%BF_ Techniques _ Source(s) Running Sprints 14-24 36 13.0 H (95, 125, 140) Middle distance 14-24 4712.8 H (95, 125] Distance 15.6 127 15.4 H 07) 25.2 63 17.1 H (28, 95, 140) 32.4 20 148 HLA (58, 139] 43.8 10 18.3 H (129) Marathon 19-24 8. Ibo H (95) 29.2 cal 16.6 A (68) 37.8 3815.5 H (126) Skiing Alpine 19.5 13 206 A 60] Nordic 17.6 15 22.7 A [109] Nordic 22.1 20173 HA (60, 108, 120) Swimming Sprint — 4 146 H {40] Middle-distance - 7 24.7 H (140) Distance. — 4 171 H (140) Synchronized 19.9 28 223 H (84, 105) All distances 17.5 63 17.3 H (83, 88, 132) Field events Discus, shot, javelin 17.8 25 23.8 H (125, 140} 19-94 ig 982) H (95) Jumps 17.4 13° 12.9 H [125] 19-94 a 17s H (95) Gymnastics 19.5 63 15.2 HA (88, 119, 121) Basketball 19.2 3620.8 H (118, 127] Mountaineering 32.7 6 187 H [39] Volleyball 20.6 48 199 H (78, 101} Lacrosse 23.0 7 23.1 A [142] Tennis 15.8 10 oe H [98] 39.0 oo 20.3 H [130] Orienteering 29.0 bs 18.7 A (74) Pentathlon 215 oo H (79] %BF = percent body fat +H = hydrostatic weighing; A = anthropometric,40 | Drinkwater women runners in their 30s have body fat levels close to the 15 percent average for young men. Only in activities where a high %BF or weight will not detract from performance do these female athletes approach levels considered average for young women [137]. Body composition values for groups drawn from the general pop- ulation are often presented as “norms,” a term often interpreted as “normal” by those unacquainted with statistical terminology. Whether 25% BF is normal for young women might be questioned. Data cited by Wilmore [140] were collected prior to 1970, with a weighted mean for 336 women in their early 20s of 25.4% BF. Data available since 1973 for 281 women of similar age average 22.4% BF (Table 5). This may be due to a sampling artifact or methodological differences, but it may also reflect the more active life-style of women in recent years. If one considers physical activity a normal part of living, perhaps “optimum” levels of %BF might be derived from body composition studies of active women of all ages and used as a guide for defining the “reasonable” bounds mentioned earlier. However, one must be careful in translating group means into criteria for individuals. There are limitations to our present techniques for assessing %BF. Wilmore [138] questions the basic assumptions underlying the densitometric method, that the density of the lean tissue is known and does not vary between individuals and that the proportion of bone and muscle comprising lean mass is a constant. Where these assumptions are violated there will be errors in the prediction of %BF for people who differ from the norm. All body composition figures should be viewed as estimates, particularly for athletes and older women, who are most likely to vary from the norm in the density of lean tissue. When facilities for hydrostatic weighing are not available, anthro- pometric methods are often used to predict %BF. It has long been recognized that prediction equations tend to be population specific. Flint et al. [50] tested 11 regression equations derived from female subjects by a number of different investigators. The validation sample ranged in age from 12 to 78 years and included both active and sedentary women. The 0.95 confidence interval for predicting %BF for the young females ranged from +5 percent to +8 percent body fat for the 11 equations. The authors concluded that the equations available in the literature were not adequate for research or clinical application. Since then Jackson et al. [67] have published a series of generalized equations using nonlinear regression equations and in- cluding age as an independent variable. While the standard error of prediction for their recommended equation is still high, ranging from 3.3 to 4.6% body fat across age groups in a cross-validation sam- ple, the concept of a generalized equation based on sound statisticalWomen and Exercise: Physiological Aspects. | 41 analyses deserves additional cross-validation reports from other in- vestigators. BLOOD PROFILES Several basic physiological differences between the sexes are found among hematological indices. Women, for example, have lower hemoglobin concentration, lower hematocrit, higher concentration of high-density lipoproteins (HDL), and higher estrogen and lower androgen levels than men. Since some of these variables are thought to be responsible for sex differences in strength and aerobic power, investigators have been examining blood profiles for other gender differences that might affect performance or that might vary as a result of strenuous exercise. Iron deficiency is one of the most frequent nutritional problems among individuals of both sexes. However, the monthly menstrual flow of the female results in an average daily loss of iron (1.3 mg) more than double that of the male (0.6 mg). If training further depletes the iron stores, women might be more at risk for iron de- ficiency anemia. Wirth et al. [141] measured serum iron concentra- tions (SeFe) in young women before and after ten weeks of training at 70% Vo, max three times a week for 20 to 25 minutes. They found no difference between the experimental and control groups in SeFe at the end of the period. However, the authors pointed out two problems with their protocol: (1) SeFe is not a measure of iron stores and may decrease only when iron stores are depleted; and (2) the training regimen may not have been of sufficient magnitude to affect iron stores. Pate et al. [93] avoided both of these problems by assessing iron stores indirectly from the percent saturation of transferrin and by examining athletes before and after their sport seasons. Of 46 women, three were iron deficient (saturation < 15%) and five borderline deficient (saturation 15-20%) before their season began. During the postseason tests, four were iron deficient and four others were bor- derline. Half the women in this study were given iron supplements (167 mg ferrous sulfate daily) throughout the season. Of the 26 athletes tested at the end of the study, eight were identified as having transferrin saturations below 20 percent. Three of these were in the iron supplementation group. Statistical analysis of the data confirmed a lack of significant effect from the iron supplements. Cooter and Mowbray [22] reported similar results for female basketball players after a four-month period of iron supplementation (18 mg ferrous fumarate 5 days per week). However, none of their athletes were iron deficient before or after the season. In at least one study [66]42 | Drinkwater where the athletes were initially iron deficient, oral supplements im- proved iron status in 96 percent of the cases. A recent study by Clement and Asmundson [20] raises the possi- bility that long-distance runners may be more at risk for iron defi- ciency than other athletes. The mean transferrin saturation for 17 women runners was 19.2 percent, compared to 24.3 percent for 35 male runners. Both groups of runners had lower values than the women athletes (X = 30.2%) described by Pate et al. [93], none of whom were runners. Clement and Asmundson [20] identified 80 percent of the women and 30 percent of the men as latent iron deficient, suggesting that women are more at risk, yet a report from Sweden of a comprehensive study of iron status in eight male distance runners found no iron or only a trace of iron in bone marrow samples from all eight [43]. Nutritional deficiencies, low absorption, and in- creased iron loss affect iron stores regardless of gender. The addi- tional loss of iron in young menstruating females adds weight to the suggestion of Pate et al. [93] that tests of iron status be included in the medical screening of female athletes. Since women do have lower resting levels of hemoglobin (Hb) and hematocrit (Hct) than men, the effect of strenuous exercise on these two hematological indices is of some concern. Puhl and Runyan [102] followed 19 women through nine weeks of aerobic, strength, and flexibility training, drawing blood for analysis at 0, 2, 5, 7, and 9 weeks. The authors report a decrease in Hb, Het, and red blood cell (RBC) count by the second week, continued low levels through the seventh week, and a return to near normal values by the ninth week. Whether the initial decrease was due to increased RBC destruction or an increase in plasma volume was not determined. However, con- current with the drop in RBC was an increase in mean corpuscular volume (MCV), which suggests selective destruction of older RBCs, rather than hemodilution. If so, there may have been a change in ‘on stores even though Hb and Hct remained within the normal range throughout the nine weeks. Wells et al. [136] examined acute hematological changes in men and women after a marathon and at 4, 8, and 24 hours later. No differences between men and women were reported, and the pattern of changes was similar for the two groups. Hct, Hb, RBC, and serum electrolytes did not vary from control values at any point. There was a marked increase in white blood cells immediately after the race that was due to an increase in polymorphonuclear neutrophils rather than in lymphocytes. Both were back to normal 24 hours later. While some of these changes differ from those reported elsewhere, possibly due to a cooler ambient temperature, there were no differences response between the sexes. Basal levels of serum constituents alsoWomen and Exercise: Physiological Aspects | 43 are similar for male and female runners. Dale et al. [29] found all values of the 20 variables measured for 22 women marathon and distance runners were within the normal range. Runners did have higher enzyme (SGOT, SGPT, and LDH) levels than sedentary con- trols, a difference also seen between male runners and nonrunners [82]. This presumably reflects the greater physical activity of the athlete. An inverse relationship between high-density lipoprotein (HDL) concentration and coronary artery disease has focused attention on the role of exercise in increasing HDL levels. A 1979 review by Wood and Haskell [143] summarized the literature dealing with the influ- ence of exercise on plasma lipids and lipoproteins for both men and women. Even though women normally have higher levels of HDL than males [104], female as well as male athletes have higher HDL concentrations than their sedentary counterparts (29, 33, 131, 143, 144]. The mean level for 71 female distance and marathon runners was 78.4 mg/100 ml; for sedentary controls it was 57.9 mg/100 ml. While distance running has received the most attention as a vehicle for increasing HDL levels, female tennis players (HDL = 73.9 mg/ 100 ml), women rowers (HDL = 65.9 mg/100 ml) and female middle- distance runners (HDL = 62.4 mg/100 ml) all have values higher than those of male athletes in the same sports and higher than the norm for sedentary women (33, 82]. For both men and women the HDL subfraction HDL, appears primarily responsible for the in- crease in total HDL [143]. COMMENT The events of the last 10 years, both scientific and athletic, have shown that women respond to strenuous training programs as men do. They improve cardiovascular fitness, increase muscle strength and endur- ance, decrease body fat, and improve performance. At the same time it appears that the current criterion for endurance capacity, VO, max, is higher for men than for women, no matter how it is expressed. This poses a problem for investigators who plan to match men and women on cardiovascular fitness for research purposes. Hopefully more investigators will become interested in resolving this problem in experimental design. Another problem related to design is the selection of sample size. Many of the studies cited in this review have small samples. When the authors report that an observed difference between groups is not significant, one is left wondering if the “power” of the statistical test was high enough to identify a real difference if indeed one did exist. Since there are methods for determining appropriate sample sizes44 | Drinkwater based on variability of the measurements, the desired level of sig- nificance, and the identification of a physiologically meaningful dif- ference, perhaps it is time for physiologists to report the power of their statistical tests as well as the levels of significance. Much of the research during the last decade has been descriptive rather than experimental. The results have been useful for docu- menting the capabilities of women in general and female athletes in particular. Studies of this nature will continue to be of interest but perhaps might be offered as brief notes rather than full-length ar- ticles. Among the experimental studies were a number that compared men and women on some variable without a hypothesis to suggest why there might be gender differences. Since so few of the older studies included women as subjects, it is understandable that inves- tigators are tempted to repeat on women all studies previously con- ducted on men. Without a reasonable hypothesis as to why sex differences might exist, the temptation should be resisted. It is refreshing to see a number of studies using male and female subjects to answer basic physiological questions. In many cases the data are analyzed without regard to gender, probably because of the small sample size in each group. While qualitative responses to ex- ercise are similar for men and women, some quantitative differences do exist. Even when these differences reflect body size, it is often useful to have the data available for each sex. Perhaps authors could provide the means and variability data for both genders, even when they are analyzed as a unit. 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