letters to nature

................................................................. along one long rutile rod (Fig. 1d). Backscattered electron images
The possible subduction of (see Supplementary Information) were used to estimate the
amounts of the exsolutions. The rods increase in concentration
continental material to with distance away from a rod-free edge zone (50–400 mm wide) to
a maximum of 4–7 vol.% for clinopyroxene, 1–2 vol.% for rutile
depths greater than 200 km and 0.5–1 vol.% for apatite in the core.
Overall, the rods occur in groups of parallel rods; each group is
Kai Ye, Bolin Cong & Danian Ye orientated in one of four directions. The effect is to show an
equilateral triangular form in cross-section (Fig. 1a). Universal-
Laboratory of Lithosphere Tectonic Evolution, Institute of Geology and Geophysics, stage measurements show that the long-axis orientations of the rods
Chinese Academy of Sciences, Beijing 100029, PO Box 9825, China are consistent with four cubic h111i directions. In a periodic
.................................. ......................... ......................... ......................... ......................... ........ cylindrical packing model13, these are dominant glide planes.
Determining the depth to which continental lithosphere can be Their concentration, topography and orientation suggest that the
subducted into the mantle at convergent plate boundaries is of rods of clinopyroxene, rutile and apatite in the porphyroblastic
importance for understanding the long-term growth of super- garnet are generated by exsolution, rather than by epitaxial replace-
continents as well as the dynamic processes that shape such ment or accidental inclusion. It is reasonable to assume that the
margins. Recent discoveries of coesite and diamond in regional cubic oxygen sublattice of garnet would have the same relation to
ultrahigh-pressure (UHP) metamorphic rocks has demonstrated the pseudohexagonal oxygen sublattice of clinopyroxene. This
that continental material can be subducted to depths of at least would imply a topotaxy between garnet and hexagonal prismatic
120 km (ref. 1), and subduction to depths of 150–300 km has been cpx in Fig. 1c as: (100)cpx // (110)grt; (010)cpx // (1̄10)grt; (1̄11)cpx //
inferred from garnet peridotites in orogenic UHP belts based on (221)grt.
several indirect observations2–5. But continental subduction to Mineral compositions were determined by means of a Camebax
such depths is difficult to trace directly in natural UHP meta- SX51 microprobe analyser. Longer counting times were pro-
morphic crustal rocks by conventional mineralogical and petro- grammed for Ti (20 s), Cr (20 s), P (60 s) and Na (60 s) in garnet
logical methods because of extensive late-stage recrystallization than for other common elements (10 s). Representative analyses are
and the lack of a suitable pressure indicator. It has been predicted listed in Table 1. The two stages of garnet and clinopyroxene differ
from experimental work, however, that solid-state dissolution of widely in their composition. The porphyroblastic garnet shows a
pyroxene should occur in garnet at depths greater than 150 km clear zonation. The exsolution-rich core contains higher Mg and Fe
(refs 6–8). Here we report the observation of high concentrations concentrations, and a lower Ca concentration. The exsolution-free
of clinopyroxene, rutile and apatite exsolutions in garnet within rim has a composition similar to that of the matrix garnet. It
eclogites from Yangkou in the Sulu UHP metamorphic belt, contains higher Ca, and lower Fe and Mg (Table 1). Particularly
China. We interpret these data as resulting from the high-pressure high concentrations of P2O5 (0.06–0.17 wt%) and Na2O (0.05–
formation of pyroxene solid solutions in subducted continental 0.06 wt%) were detected in the core of the porphyroblastic garnet.
material. Appropriate conditions for the Na2O concentrations and However, the Ca-rich rim and the matrix garnet contain almost no
octahedral silicon observed in these samples are met at depths P2O5 (, 0.03 wt%) or Na2O (, 0.02 wt%) (Table 1). The exsolved
greater than 200 km. cpx-I contains lower jadeite (Xjd = 0.17–0.25) than the matrix
Supercontinents are generally thought to grow through subduc- omphacite (Xjd = 0.29–0.35). (Here Xjd is the volume per cent of
tion and collision of smaller continental blocks—as is happening in jadeite component.) The average composition of the exsolved cpx-I,
north India and southwest China, where the Indian continental
plate is subducting below the Eurasian continent, and causing the
uplift of the Tibetan plateau. Subduction of the South China
continent under the North China continent in the Triassic9 period a grt-I b
consolidated the old Eurasia continent, and resulted in the forma- cpx
tion of various coesite- and diamond-bearing UHP metamorphic
rocks in the oregon in eastern China (for a review, see ref. 10). The
Sulu UHP metamorphic belt is the most deeply exhumed part of the rut
Dabieshan-Sulu oregon. This provides the possibility of tracing the
subduction depths of continental materials from fresh samples in
the Sulu UHP belt. grt-II+cpx-II+rut-II+ap-II
Unusual eclogites were found as lenses in a garnet-peridotite layer c d
intercalated with layers of UHP metamorphosed crustal igneous cpx
composite11 at Yangkou. Partial transformation from low-pressure ap
igneous textures (plagioclase stable) to UHP textures (coesite
stable) have been reported in some meta-gabbroic layers of the
composite12. Unlike the common eclogites with near equigranular
rut
texture in the world-wide UHP belts, these eclogites contain an
rut
older generation of garnet (grt-I) which occurs as larger (up to
2 cm), purple, plastically deformed porphyroblast in the matrix of
later medium-grained (0.05–0.5 mm) garnet (grt-II), omphacite Figure 1 Transmitted light photomicrographs of garnet (grt) with exsolution rods of
(cpx-II), rutile (rut-II) and apatite (ap-II) (Fig. 1a). Grt-I exhibits clinopyroxene (cpx), rutile (rut) and apatite (ap) along crystallographically controlled
abundant fine-grained euhedral rods or needles of clinopyroxene planes. a, Porphyroblastic garnet (grt-I) in the matrix of garnet (grt-II), omphacite (cpx-II),
(cpx-I), rutile (rut-I) and apatite (ap-I) (Fig. 1a). The clinopyroxene rutile (rut-II) and apatite (ap-II), the exsolution rods oriented in an equilateral triangular
rods are spindle-shaped in the long axis section (Fig. 1b), and form. Scale bar, 1 mm. b, Spindle-shaped long-axis section of clinopyroxene rods in
hexagonal prismatic in the basal section (Fig. 1c). Apatite rods occur porphyroblastic garnet. Scale bar, 0.1 mm. c, Hexagonal prismatic basal sections of
as hexagonal prism or rhombohedra. Like the hexagonal prismatic clinopyroxene rods in porphyroblastic garnet. Note that each group of pyroxene crystal
clinopyroxene, each group of apatite crystal planes is parallel to one planes is parallel to one group of needle-like rutile rods. Scale bar, 0.06 mm. d, Three
group of rutile rods. In cases, several short apatite rods are dotted short apatite rods dot one long rutile rod. Scale bar, 0.1 mm.

734 © 2000 Macmillan Magazines Ltd NATURE | VOL 407 | 12 OCTOBER 2000 | www.nature.com
 letters to nature
rut-I and ap-I were restored to the core composition of grt-I bimineralic eclogitic assemblage, the pressure conditions under
following the estimated volume proportions of grt91.5cpx7rut1ap0.5. which the garnet was once homogeneously pyroxene-substituted
The calculated original garnet (no. 6 in Table 1) is depleted in Al must be inferred from experimental data on basaltic systems under
(with respect to the chemical formula) and contains 0.04 more Si subduction conditions. As pressure increases, the solubility of
than the ideal value of 3 for 12 oxygens, necessitating the existence pyroxene in eclogitic garnet increases gradually7,20, and the Na, Ti
of 0.06 (SiTotal + PIV − 3.00) 6-fold Si in octahedral coordination and P contents in garnet also have a positive pressure dependence18.
(Table 1). Here the roman numerals indicate the degree of coordi- The calculated Na2O content (0.34 wt%) and octahedral silicon
nation of the atoms. It also exhibits high Na2O (0.34 wt%), TiO2 values (0.06) of the original Yangkou garnet are higher than those of
(1.07 wt%) and P2O5 (0.28 wt%) values. A slightly lower average the garnets synthesized21 at 7 GPa and 1,000 8C using starting
Na2O content (0.28 wt%) was confirmed for the core of the material of a natural eclogite from Dabieshan, and also higher
porphyroblastic garnet with the highest amount of clinopyroxene than those of garnet synthesized20 at 7 GPa and 1,400 8C in a MORB
exsolution by broad-beam electron microprobe analysis. system: but the values of 0.34 wt% and 0.06 are consistent with
Experiments have predicted the structural transformation of garnet synthesized7 at 7.6 GPa and 1,200 8C in an olivine tholeiite
pyroxene to garnet and the solid-state dissolution of pyroxene system. In these synthesis experiments, the temperatures of 1,400 8C
into garnet at very-high-pressure conditions (5–17 GPa)6–8. Con- and 1,200 8C are higher than in the subduction regime, and the
version of pyroxene to the garnet structure requires some of the Na2O content in garnet is known20 to be higher at higher tempera-
four-fold tetrahedral Si in pyroxene to be changed to six-fold tures. Thus, it is reasonable to conclude that the original garnet in
octahedral coordination in converted majoritic garnet, because the Yangkou eclogites was formed at pressures greater than 7 GPa.
the ratio of Si to O in pyroxene is 4:12, rather than the garnet This puts the origin of the studied Yangkou eclogites at depths
value of 3:12. From a crystal chemistry point of view, substitution of greater than 200 km.
NaVIII + SiVI → CaVIII + AlVI in the garnet structure can describe The tectonic settings of the protolith of Yangkou eclogite and the
these transformations14. Direct access to xenoliths sampled by host garnet-peridotite are not clear. However, lower-pressure
diamond kimberlite pipes from the stable mantle lithosphere at mineral inclusions have been reported in garnet of similar garnet-
depths greater than 300 km have revealed pyroxene dissolution in peridotite from nearby localities in the same belt22. This implies
homogeneous garnet included in diamond15, and decompression possible crustal association of the studied eclogite before subduc-
resulted in exsolution of clinopyroxene in garnet16, which is iden- tion. An alternative interpretation is that the peridotite was under-
tical to the occurrence of clinopyroxene in garnet reported here. plated at the base of the continental basement then underwent
Experiments17,18 have shown that garnet can host high Ti, Na and P ultrahigh-pressure metamorphism22. But regardless of the exact
contents under very-high-pressure conditions. Phosphorus has tectonic settings, our results have implications for the dynamics
been suggested18 to enter the garnet structure through the coupled of continental subduction and exhumations of the subducted rocks.
substitution of PIV + NaVIII → SiIV+ CaVIII. Ringwood and co- Density calculations based on experimental data confirm that
workers6,17 proposed that (CaNa2)Ti2Si3O12 and (Ca2Na)(AlTi)- subducted oceanic slab is substantially denser than the surrounding
Si3O12 are probable Ti-bearing garnet components, based on com- pyrolitic mantle at depths of 100–650 km (ref. 7). Oceanic slab has
positions of garnets synthesized under high-pressure conditions, been observed to be subducted to depths exceeding 600 km in the
and they hence suggested that Ti enter garnet through substitution mantle23. Subduction of the less-dense continental crust to great
of NaVIII + TiVI → CaVIII + AlVI. Such Na-, P- and Ti-bearing garnets mantle depths is traditionally thought to be difficult, because the
had been reported in ultra-deep xenoliths in diamond kimberlite operation of buoyancy forces would drive it back to the crustal level.
pipes14,15 and UHP metamorphic crustal rocks19. However, subducted continental crust, and terrigenous and pelagic
Because of the lack of a suitable pressure indicator for the sediments, can be important in the geochemical evolution of the

Table 1 Representative compositions of clinopyroxene (cpx) and garnet (grt)

1 2 3 4 5 6 7 8 9
...................................................................................................................................................................................................................................................................................................................................................................
Mineral cpx cpx grt grt grt grt grt grt grt
Group E M CP RP M O SY SY SY
SiO2 55.36 55.56 39.39 38.93 39.21 40.32 41.43 41.49 40.94
TiO2 0.06 0.01 0.11 0.06 0.06 1.07 0.15 0.03 1.49
Al2O3 6.14 7.90 21.37 21.01 20.87 20.18 22.46 22.77 22.56
Cr2O3 0.01 0.06 0.04 0.00 0.05 0.06 – 0.19 –
FeO 3.85 4.03 21.75 18.60 18.92 20.19 13.61 13.05 12.29
MgO 12.50 10.82 9.57 6.49 5.98 9.41 12.28 16.20 9.96
MnO 0.05 0.05 0.49 0.43 0.45 0.47 – 0.26 –
CaO 17.51 16.14 7.12 14.45 14.90 8.36 9.67 5.61 12.46
Na2O 4.31 5.54 0.06 0.01 0.02 0.34 0.38 0.06 0.27
K2O 0.00 0.01 0.00 0.00 0.00 0.00 – 0.06 –
P2O5 0.01 0.00 0.09 0.01 0.00 0.28 – – –
Total 99.78 100.12 99.99 99.99 100.46 100.68 99.98 99.72 99.97
...................................................................................................................................................................................................................................................................................................................................................................
Cation proportions based on 6 (for cpx) and 12 (for grt) oxygens
Si 1.995 1.992 3.001 2.991 3.005 3.039 3.046 3.019 3.019
Ti 0.002 0.000 0.006 0.003 0.003 0.061 0.008 0.002 0.083
Al 0.261 0.334 1.919 1.902 1.885 1.793 1.946 1.953 1.961
Cr 0.000 0.002 0.002 0.000 0.003 0.004 – 0.011 –
Fe 0.116 0.121 1.386 1.195 1.213 1.273 0.837 0.794 0.758
Mg 0.671 0.578 1.087 0.743 0.683 1.057 1.346 1.757 1.095
Mn 0.001 0.002 0.032 0.028 0.029 0.030 – 0.016 –
Ca 0.676 0.620 0.581 1.189 1.224 0.675 0.762 0.437 0.984
Na 0.301 0.385 0.009 0.001 0.003 0.050 0.054 0.008 0.039
K 0.000 0.000 0.000 0.000 0.000 0.000 – 0.006 –
P 0.000 0.000 0.006 0.001 0.000 0.018 – – –
...................................................................................................................................................................................................................................................................................................................................................................
Values given are for analyses of minerals in Yangkou eclogites (nos 1-5); no. 6 represents calculated original garnet; nos 7, 8 and 9 are garnets synthesized in basaltic systems at 7.6 GPa and 1,200 8C by
Irifune et al.7, 7 GPa and 1,000 8C by Okamoto and Maruyama21, and 7 GPa and 1,400 8C by Ono and Yasuda20, respectively. Abbreviations; cpx: clinopyroxene; grt: garnet; E: exsolution; M: matrix, CP:
core of porphyroblastic garnet, RP: rim of porphyroblastic garnet, O: original garnet, SY: synthesized garnet.

NATURE | VOL 407 | 12 OCTOBER 2000 | www.nature.com © 2000 Macmillan Magazines Ltd 735
letters to nature
source of many alkali intraplate hotspot magmas—such sources are
at extreme depths in the mantle24–26. Recent discoveries of coesite
and diamond in eclogites, metamorphic sedimentary rocks1 and
even granitic gneisses27 in continental subduction orogenic belts
reveal that large, coherent volumes of continental materials could be
subducted to depths of more than 120 km. The present study
further extends continental subduction to depths of more than
200 km. Experiments and density calculations8 show that the
densities of the continental crust are substantially smaller than
that of the surrounding pyrolitic mantle (3.4 g cm−3) at pressures
lower than 6 GPa (3.0 g cm−3), slightly less-dense in the interval of
6.5–8.5 GPa (3.2–3.3 g cm−3), and much denser above 9–10 GPa
until 24 GPa (3.7–3.9 g cm−3)8. Once the subducted continental
materials reach a depth of about 200 km, their densities approach
that of the surrounding mantle, and exceed the density of the
surrounding mantle below 300 km. Thus, the buoyancy forces are
effectively cancelled and entrainment by the sinking slab becomes
the predominant influence. In the Sulu case, a subducted oceanic
slab is thought to have hauled the following continental materials to
depths greater than 200 km. Only one mechanism is reasonable
for exhumation of such deeply subducted rocks: that they are rafted
by rocks beneath them, propelled by buoyancy forces2. Thus, a
detachment was required along the subducting slab at some depths
between 200 and 300 km, so that the buoyancy forces could drive
the deeply subducted continental materials back to the crustal
level.
Much larger-scale extremely deeply subducted crustal materials
may be present in the Sulu UHP belt. Such materials would be
indicated by the presence of appropriate mineralogical or micro-
textural evidence—such as TiO2 with the a-PbO2 structure (stable
at 5–7 GPa), K-wadeite (6.5–7 GPa), or high-pressure C2/C clin-
oenstatite (6–7 GPa). We note that the presence of high-pressure
C2/C clinoenstatite was previously predicted28 for the Alpe Arami
peridotite, and that this was subsequently confirmed3. M specialization in evolving
Received 13 June; accepted 24 August 2000.
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96–131 (Cambridge Univ. Press, New York, 1995).
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than 300 kilometers. Science 271, 1841–1845 (1996).
3. Bozhilov, K. N., Green, H. W. & Dobrzhinetskaya, L. Clinoenstatite in Alpe Arami peridotite:
additional evidence of very high pressure. Science 284, 128–132 (1999).
4. Van Roermund, H. L. M. & Drury, M. R. An ultra-deep (.200 km) orogenic peridotite body in
Western Norway. Eos 79, F971 (1998).
5. Yang, J. J., Godard, G., Kienast, J. R., Lu, Y. & Sun, J. Ultrahigh-pressure (60 kbar) magnesite-bearing
garnet peridotites from northeastern Jiangsu, China. J. Geol. 101, 541–554 (1993).
6. Ringwood, A. E. & Major, A. Synthesis of majorite and other high pressure garnets and perovskites.
Earth Planet. Sci. Lett. 12, 411–418 (1971).
7. Irifune, T., Sekine, T., Ringwood, A. E. & Hibberson, W. O. The eclogite-garnet transformation at high
pressure and some geographical implications. Earth Planet. Sci. Lett. 77, 245–256 (1986).
8. Irifune, T., Ringwood, A. E. & Hibberson, W. O. Subduction of continental crust and terrigenous and
pelagic sediments: an experimental study. Earth Planet. Sci. Lett. 126, 351–386 (1994).
9. Ames, L., Tilton, G. R. & Zhou, G. Z. Timing of collision of the Sino-Korean and Yangtze cratons:
U-Pb zircon dating of coesite-bearing eclogites. Geology 21, 339–342 (1993).
10. Cong, B. L. Ultrahigh-Pressure Metamorphic Rocks in the Dabieshan-Sulu Region of China (Science
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11. Wallis, S. R. et al. Occurrence and field relationships of ultrahigh-pressure metagranitoid and coesite
eclogite in the Su-Lu terrane, eastern China. J. Geol. Soc. Lond. 154, 45–54 (1997).
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the Sulu terrane, eastern China. J. Metamorph. Geol. 15, 183–202 (1997).
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Dora Maira Massif, Western Alps: detailed petrography, mineral chemistry and P-T path. Contrib.
Mineral. Petrol. 108, 1–21 (1991).
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20. Ono, S. & Yasuda, A. Compositional change of majoritic garnet in a MORB composition from 7 to 17
GPa and 1400 to 1600 oC. Physics Earth Planet. Inter. 96, 171–179 (1996).
21. Okamoto, K. & Maruyama, S. Multi-anvil re-equilibration experiments of a Dabie Shan ultrahigh-
pressure eclogite within the diamond-stability fields. Island Arc 7, 52–69 (1998).
22. Zhang, R. Y., Liu, J. G. & Cong, B. Petrogenesis of garnet-bearing ultramafic rocks and associated
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186 (1995).
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Supplementary Information is available on Nature’s World-Wide Web site
(http://www.nature.com) or as paper copy from the London editorial office of Nature.
Acknowledgements
We thank P. Xu for programming microprobe analyses, and H. W. Green for comments
that significantly improved this Letter. This work was supported by the National Natural
Science Foundation of China
Correspondence and requests for materials should be addressed to K.Y.
(e-mail: yekai@hotmail.com).
.................................................................
The population genetics of ecological
Escherichia coli populations
Vaughn S. Cooper & Richard E. Lenski
Center for Microbial Ecology, Michigan State University, East Lansing,
Michigan 48824, USA
.......................................... ......................... ......................... ......................... .........................
When organisms adapt genetically to one environment, they may
lose fitness in other environments1–4. Two distinct population
genetic processes can produce ecological specialization—muta-
tion accumulation and antagonistic pleiotropy5–8. In mutation
accumulation, mutations become fixed by genetic drift in genes
that are not maintained by selection; adaptation to one environ-
ment and loss of adaptation to another are caused by different
mutations. Antagonistic pleiotropy arises from trade-offs, such
that the same mutations that are beneficial in one environment
are detrimental in another. In general, it is difficult to distinguish
between these processes5–8. We analysed the decay of unused
catabolic functions in 12 lines of Escherichia coli propagated on
glucose for 20,000 generations9,10. During that time, several lines
evolved high mutation rates11. If mutation accumulation is impor-
tant, their unused functions should decay more than the other
lines, but no significant difference was observed. Moreover, most
catabolic losses occurred early in the experiment when beneficial
mutations were being rapidly fixed, a pattern predicted by
antagonistic pleiotropy. Thus, antagonistic pleiotropy appears
more important than mutation accumulation for the decay of
unused catabolic functions in these populations.
We founded 12 populations from a strain of E. coli, and they all
adapted to minimal medium supplemented with glucose9,10.
Competitive fitness increased rapidly in the first few thousand
generations. Fitness continued to improve, but the average rate of
NATURE | VOL 407 | 12 OCTOBER 2000 | www.nature.com
Supplementary Figure Caption:

Backscattered electron imagine showing high concentration of clinopyroxene (dark grey) rods in
garnet.