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We showed that the new metabolic rate relation13 can be directly linked to the total energy consumed in a lifespan if a constant number ({mathrm{N}}_{mathrm{r}}) of respiration cycles per lifespan is conjectured, and a corrected relation for the total energy consumed in a lifespan was found [Eq. (1)] that can explain the origin of variations in the ‘rate of living’ theory2,5 and unify them into a single formulation. It is important to note that Eq. (1) is a direct consequence of combining the two empirical relations mentioned (the new metabolic rate relation and the relation of the total number ({mathrm{N}}_{mathrm{r}}) of respiration cycles per lifespan) and is not an assumption (based on the lifespan energy expenditure per gram) as in the traditional ‘rate of living’ theory2,5. We test the validity and accuracy of the predicted relation [Eq. (1)] for the total energy consumed in a lifespan with (sim 300) species representing different classes of living organisms, and we find that the relation has an average scatter of only 0.3 dex, with 95% of the organisms having departures of less than a factor of (pi) from the relation, despite the difference of (sim 20) orders of magnitude in body mass.Successful testing of predictions is crucial in any proposed theory according to Popper’s deductive method of falsification (27), which is the criterion for identifying a successful scientific theory. Therefore, the success of the predicted Eq. (1) that is displayed in Fig. 1 implies that the corrected metabolic rate relation13 has passed an initial test. This prediction also reduces any possible interclass variation in the relation, which has been considered the most persuasive evidence against the ‘rate of living’ theory, to only a geometrical factor and strongly supports the conjectured invariant number ({mathrm{N}}_{mathrm{r}} sim 10^8) of respiration cycles per lifespan in all living organisms.Invariant quantities in physics traditionally reflect fundamental underlying constraints, a principle that has also been applied recently to life sciences such as ecology21,22. Figure 2 indicates the fact that, for a given temperature, the total lifespan energy consumption per gram per ‘generalized beat’ (({mathrm{N}}_{{mathrm{b}}}^{mathrm{G}} equiv mathrm{a} {mathrm{N}}_{{mathrm{r}}} = {mathrm{a}} ,1.62 times 10^8)) is remarkably constant (around ({mathrm{E}}_{2019})), a result that is also in agreement with previous expectations based on (lifespan) basal oxygen consumption at the molecular level38. This supports the idea that the overall energetics during the lifespan are the same for all the organisms studied, as it is predetermined by the basic energetics of respiration, and therefore, Rubner’s original picture is shown to be valid without systematic exceptions but in a more general form. Moreover, since the value determined from Fig. 2 is remarkably similar to ({mathrm{E}}_{2019} {mathrm{N}}_{mathrm{r}}), it can be considered an independent determination of ({mathrm{E}}_{2019}), suggesting that ({mathrm{E}}_{2019}) is a candidate for being a universal constant and not just a fitting parameter from the corrected metabolic relation13.In addition, we showed here that the invariant total lifespan energy consumption per gram per ‘generalized beat’ comes directly from the existence of another invariant, the approximately constant total number ({mathrm{N}}_{mathrm{r}} sim 10^8) of respiration cycles per lifetime, effectively converting the ‘generalized beat’ into the characteristic clock during the lifespan. Therefore, the exact physical relation between (oxidative) free radical damage and the origin of aging is most likely related to the striking existence of a constant total number of respiration cycles ({mathrm{N}}_{{mathrm{r}}}) over the lifetime of all organisms, which predetermines the extension of life. Moreover, the relation ({mathrm{t}}_{{mathrm{life}}} = mathrm{N}_{mathrm{r}}/mathrm{f}_{{{mathrm{resp}}}}) quantifies the ideas of oxidative damage by the respiratory metabolism, which are motivated mainly by biomedical considerations, into a simple mathematical form that could be included in a broader life-history framework; this is needed to produce testable predictions for the ‘free-radical’ hypothesis in the life-history context28. Future theoretical and experimental studies that investigate the exact link between the constant number ({mathrm{N}}_{mathrm{r}} sim 10^8) of respiration cycles per lifespan and the production rates of free radicals (or alternatively, other byproducts of metabolism) should shed light on the origin of aging and the physical cause of natural mortality.Although this relation ({mathrm{t}}_{{mathrm{life}}} = mathrm{N}_{mathrm{r}}/mathrm{f}_{{mathrm{resp}}}) has only been empirically examined for mammalian vertebrates, in terms of heartbeats per lifetime, there is evidence to believe that the relative constancy of the number of respiration cycles per lifetime is more widely distributed in the animal kingdom. For example, a reptile such as the Galapagos tortoise with a life expectancy of 177 years and a respiration rate of 3 breaths/min has (2.8 times 10^8) breaths per lifetime29, which is within a factor of 2 of the value determined for mammals. A more different case is that of birds, which have more heartbeats/lifetime by a factor of 330; this difference is reduced to a factor of 1.5 in terms of breaths/lifetime ((mathrm{N}_{mathrm{r}} = mathrm{N}_{mathrm{b}}/{mathrm{a}}), with (hbox {a}=9) for birds and 4.5 for mammals; 17). Among fish, the average number of heartbeats/lifetime tends to be an order of magnitude less than that in mammals ((mathrm{N}_{mathrm{b}} = 7.3 times 10^8);16), for example, (mathrm{N}_{mathrm{b}} = 6.7 times 10^7) for trout31, but in such cases, the parameter a can be as low as 0.5 (i.e., a heart frequency lower than the respiratory frequency; 32), again implying a similar ({mathrm{N}}_{mathrm{r}} ,(= mathrm{N}_{mathrm{b}}/{mathrm{a}} = 1.3 times 10^8)). A more extreme difference in terms of heartbeats is the tiny Daphnia, which uses up to (1.7 times 10^7) heartbeats (at 25 C) in a short lifespan of 30 days33. Simple invertebrates, such as Daphnia, do not have a complex respiratory system with lungs and obtain oxygen for respiration through diffusion, but a “breath frequency” can be estimated from its respiration rate ((sim mu {mathrm{l}} {mathrm{O}}_2 hbox {hr}^{-1});34) divided by ({mathrm{E}}_{2019} M) (with ({mathrm{M}} sim 100 mu {mathrm{g}});35), giving ({mathrm{N}}_{mathrm{r}} = 1.5 times 10^8) respiration cycles per lifetime. In summary, a difference of two orders of magnitude in total heartbeats (between Daphnia and birds) is reduced to less than a factor of 2 in breaths per lifetime, further supporting that all organisms seem to live for the same span in units of respiration cycles (({mathrm{N}}_{mathrm{r}} sim 10^8)).It has also been suggested that an analogous invariant originates at the molecular level23, the number of ATP turnovers of the molecular respiratory complexes per cell in a lifetime, which, from an energy conservation model that extends metabolism to intracellular levels, is estimated to be (sim 1.5 times 10^{16})23. A similar number can be determined by taking into account that human cells require the synthase of approximately 100 moles of ATP daily, equivalent to (7 times 10^{20}) molecules per second. For (sim 3 times 10^{13}) cells in the human body and for a respiration rate of 15 breaths per minute, this gives (sim 9 times 10^{7}) ATP molecules synthesized per cell per breath, which for the invariant total number ({mathrm{N}}_{mathrm{r}}) of respiration cycles per lifetime found in this work, rises to the same number of (sim 1.5 times 10^{16}) ATP turnovers in a lifetime per cell, showing the equivalence between both invariants and linking ({mathrm{N}}_{mathrm{r}}) to the energetics of respiratory complexes at the cellular level.The excellent agreement between the predicted relation [Eq. (1)] and the data across all types of organisms emphasizes the fact that lifespan indeed depends on multiple factors (B, a, M, T & (mathrm{T}_{mathrm{a}})) and strongly supports the methodology presented in this work of multifactorial testing, as shown in Fig. 1, since quantities in life sciences generally suffer from a confounding variable problem. An example of this problem, illustrated by individually testing each of the relevant factors, is given in24, which for a large (and noisy) sample test for ({mathrm{t}}_{{{mathrm{life}}}} propto 1/B) shows no clear correlation. From Eq. (1), it is clear that in an uncontrolled experiment, the dependence on the rest of the parameters (M, a, T, & ({mathrm{T}}_{mathrm{a}})) might eliminate the dependence on the metabolic rate B (in fact, this may be for the same reason that Rubner’s work7 focused on the mass-specific metabolic rate B/M instead of B). This work24 finds only a residual inverse dependence of ({mathrm{t}}_{{mathrm{life}}}) on the ambient temperature ({mathrm{T}}_{{mathrm{a}}}) for ectotherms, which is expected according to Eq. (1) (Big (mathrm{t}_{{mathrm{life}}} propto {mathrm{exp}}Big ({small frac{mathrm{E}_{mathrm{a}}}{mathrm{k} {mathrm{T}}_{mathrm{a}}}}Big ) Big )).Finally, the empirical support in favor of Eq. (1) allows us to perform several estimations regarding how much the energy consumption will vary with changing physical conditions on Earth. For example, computing by how much the energy consumption will vary in biomass performing aerobic respiration as the Earth’s temperature increases is relevant in the current context of possible global warming. This is given by the factor ({mathrm{exp}}Big [{small frac{mathrm{E}_{mathrm{a}}}{{mathrm{k}}} Big (frac{1}{ {mathrm{T}}}} – {small frac{1}{ {mathrm{T}}+1}}Big ) Big ]), which for an activation energy of ({mathrm{E}}_{mathrm{a}} = 0.63 ,hbox {eV}) and a temperature of (30^{circ }hbox {C}) implies an increase of 8.3% in energy consumption per 1 degree increase in the average Earth temperature. This result can be straightforwardly applied in ectotherms since their body temperatures adapt to the environmental temperature (({mathrm{T}}={mathrm{T}}_{mathrm{a}})), but its implications for endothermic organisms are less clear. Another relevant estimation is to compute by how much B({mathrm{t}}_{{mathrm{life}}})/M would vary from Eq. (1) (i.e., the difference between Figs. 2 and 3) as a function of body temperature (T) and the ratio of heart rate to respiratory rate ((mathrm{a}= mathrm{f}_{mathrm{H}}/ {mathrm{f}}_{{mathrm{resp}}})). Variations in B({mathrm{t}}_{{mathrm{life}}})/M are relevant since this is a key quantity in the estimation of the energy allocation to fitness, which aims to explain in terms of trade offs the so-called ‘Equal Fitness Paradigm’39 that concerns why most organisms in the biosphere are more or less equally fit, other than the diversity seen in the size, form and function of living organisms on Earth.In the near future, our plan is to generate a (metabolic) theory starting from the new metabolic rate relation13 by assuming that it is the controlling rate in ecology in order to explain a variety of ecological phenomena in a similar fashion as the metabolic theory of ecology18 does using Kleiber’s law. A first step in this direction looks very promising40, as it can show that ontogenetic growth can be described by a universal growth curve without the aid of fitting parameters, can explain the origin of several ‘Life History Invariants’21 and can show how the heart rate may actually set several biological times (i.e., lifespan and generation time) and even some ecological rates (i.e., The Malthusian parameter). More

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Phylogenetic analysesA total of 21 sequences were newly generated and deposited in GenBank (Supplementary Table 1). The concatenated sequence alignment of the three loci comprised 100 sequences (38 for ITS, 30 for rpb2 and 32 for mtSSU) from 43 collections (Supplementary Table 1). The alignment was 2,004 characters long, including gaps. Multi-locus trees generated from ML and BI analyses showed similar topologies without any supported topological conflict. The multi-locus phylogeny (Fig. 1) confirmed placement of all Thai collections within the well-supported R. subsect. Amoeninae (ML = 99, BI = 1.0). Five collections from northeastern Thailand and two collections from northern Thailand form two strongly supported clades and are described below as the new species R. bellissima sp. nov. and R. luteonana sp. nov. The new species are not resolved as sister. The first species, R. bellissima, is strongly supported as sister to a clade of Australian sequestrate species that includes R. variispora T. Lebel and an undescribed Russula sp. labeled as Macowanites sp. The Indian species R. intervenosa S. Paloi, A.K. Dutta & K. Acharya is placed as sister to them with bootstrap support of 77. The second species, R. luteonana, is placed with moderate support as sister to the sequestrate European species R. andaluciana T.F. Elliott & Trappe.Figure 1ML phylogenetic tree inferred from the three-gene dataset (ITS, rpb2, mtSSU) of Russula subsection Amoeninae species, using ML and BI analyses. Three members of R. subg. Heterophyllidiae are used as outgroup. Species in boldface are new species in this study. Bootstrap support values (BS ≥ 50%) and posterior probabilities (PP ≥ 0.90) are shown at the supported branches.Full size imageThe ITS tree (Fig. 2) shows a similar topology and relationships for the studied specimens. In addition, R. intervenosa received good support (ML = 84, BI = 0.99) as sister to the clade of R. bellissima and R. variispora. Five additional ITS sequences that are grouped with strong support within R. bellissima species clade were recovered, three from Thailand, one from Laos, and one from Singapore. We did not recover any other Amoeninae ITS sequences from Thailand.Figure 2ML phylogenetic tree inferred from the ITS region of Russula subsection Amoeninae species and allied groups, using ML and BI methods. Samples in boldface are new species in this study. Bootstrap support values (BS ≥ 50%) and posterior probabilities (PP ≥ 0.90) are shown at the supported branches.Full size imageTaxonomy
Russula bellissima Manz & F. Hampe sp. nov.
Mycobank: MB 840549Holotype THAILAND, Theong district, Chiang Rai, 19°36′45”N 100°4′00”E, alt. 500 m, dry dipterocarpus forest in small groups on loamy soil, 12 July 2012, F. Hampe (Holotype: GENT FH 12-127; Isotype: MFLU12-0619).Etymology ’bellus’ = latin for beautiful, pretty, lovely; ’bellissima’ = the most beautiful. Resembling the species Russula bella which is also belonging to Russula subsection Amoeninae.Diagnosis Pileus small to medium-sized; cuticle dry, smooth, matt and pruinose, red; stipe white or with a red flush; spore ornamentation of moderately distant to dense amyloid spines or warts, frequently fused into short crests or even long wings; suprahilar spot inamyloid; hymenial cystidia and pileocystidia absent.Pileus (Fig. 3) small to medium sized, 10–50 mm diam., young hemispherical or convex, becoming plane and depressed at the centre; margin first even, when old distinctly tuberculate-striate up to 10 mm from the margin, often radially cracking; cuticle hardly peeling, radially disrupted into small patches, pruinose when young, later dry, smooth, matt and pruinose in the centre, colour near the margin when young varnish red (9C8), later red to coral red (9B6-7); near the centre deep red, blood red, dark red (10C7-8), raspberry red (10D7), strawberry red (10D8) or purple brown (10E-F8). Lamellae: 3–5 mm deep, thin, moderately dense, 6–8 at 1 cm near the pileus margin, adnexed, white, slightly anastomosing at the base; lamellulae absent, occasionally forked near the stipe; edges concolorous, entire but pruinose under lens. Stipe: 10–30 × 3–7 mm, usually narrowed towards the base, sometimes cylindrical, surface smooth, white and mainly with a distinct pastel red to red flush, occasionally completely white or sometimes also almost completely red, interior stuffed. Context: white, fragile, unchanging when damaged, reaction with guaiac after 5 s negative on both stipe and lamellae surfaces, reaction to FeSO4 and sulfovanillin negative; taste mild; odour inconspicuous. Spore print: not observed.Figure 3Basidiomata of Russula bellissima. (A) FH12-127 (Holotype). (B) FH12-158. Scale bar = 1 cm. Photos by Felix Hampe.Full size imageSpores (Figs. 4, 5) (6.9–)7.3–7.8–8.3(–8.9) × (6.1–)6.8–7.2–7.6(–8.4) µm, subglobose to broadly ellipsoid, Q = 1.01–1.1–1.2(–1.29); ornamentation of moderately distant [(4–)5–6(–7) in a 3 µm diam. circle] amyloid spines or warts, (1.1–)1.2–1.4–1.6(–1.7) µm high, fused or connected by fine line connections into often long crests or wings, [(0–)1–3(–4) fusions and the same number of line connections in a 3 µm diam. circle], crests and wings frequently branched and occasionally form closed loops, isolated elements dispersed, edge of crests and wings irregularly wavy; suprahilar spot moderately large, inamyloid. Basidia: (30.5–)34.5–44.1–53.5(–65.0) × (10.5–)11.5–12.6–14.0(–16.0) µm, broadly clavate or obpyriform, 4-spored; basidiola cylindrical, ellipsoid or broadly clavate, ca. 5–10 µm wide. Hymenial cystidia on lamellae sides: absent. Lamellae edges: covered by densely arranged or fasciculate marginal cells. Marginal cells: (27.0–)38.5–46.4–54.5(–61.0) × (5.0–)5.5–6.7–7.5(–9.0) µm; subulate or narrowly lageniform, apically attenuated and constricted to ca. 1–2 µm, sometimes slightly moniliform or flexuous. Pileipellis: (Fig. 6) orthochromatic in Cresyl Blue, gradually passing to the underlying context, 200–300 µm deep; suprapellis 60–130 µm deep, composed of erect or ascending hyphal terminations forming a dry trichoderm, well delimited from 140 to 210 µm deep subpellis composed of horizontally oriented, strongly gelatinized narrow hyphae. Subpellis not well delimited from the underlying context, elongate hyphae gradually changing to sphaerocytes. Acid- resistant incrustations: absent. Hyphal terminations near the pileus margin: composed of long apically attenuated terminal cell and a chain of 1–4 ovoid to barrel shaped, short unbranched cells with one distinctly longer apical cell; constricted on septa, usually not flexuous, oriented towards the pileus surface, usually thin-walled, sometimes slightly thick-walled (up to 1 µm thick); terminal cells mainly subulate or lageniform, apically attenuated and acute, measuring (19–)27.5–38.3–49.0(–66.5) × (3.3–)4.5–5.8–7.0(–9.0) µm, rarely with a forked apex, mixed with dispersed, cylindrical or ellipsoid, distinctly shorter, obtuse terminal cells measuring (7.5–)11.5–17.8–29.5(–42.5) × (3.0–)4.0–4.5–5.0 µm; subterminal cells measuring (4.5–)5.5–8.3–11.5(–16.0) × 4.5–5.3–6.0(–7.0) µm. Hyphal terminations near the pileus centre: similar in shape and also with a mixture of long acute and short obtuse terminal cells, acute ones measuring (12.0–)22.0–35.2–48.5(–79.0) × (2.5–)3.5–4.9–6.5(–8.0) µm, obtuse ones more frequent, measuring (6.5–)8.5–12.0–15.5(–22.0) × (3.5–)4.0–4.9–6.0(–7.5) µm. Primordial hyphae or pileocystidia: absent. Cystidioid hyphae and oleipherous hyphae not observed.Figure 4Hymenial elements of Russula bellissima (holotype, FH 12-127). (A) Basidia and basidiolae. (B) Marginal cells. (C) Spores as seen in Melzer’s reagent. Scale bar = 10 µm, but only 5 µm for spores.Full size imageFigure 5Scanning electron microscope photo of spore ornamentation. Russula bellissima (holotype, FH 12-127). Scale bar = 2 μm.Full size imageFigure 6Elements of the pileipellis of Russula bellissima (holotype, FH 12-127). (A) Hyphal terminations near the pileus margin. (B) Hyphal terminations near the pileus centre. Scale bar = 10 μm.Full size imageAdditional material studied THAILAND, Chiang Mai Province, Mae On District, about 3 km from Tharnthong lodges, 18° 51′ 55″ N 99° 17′ 23″ E, alt. 725 m, Dipterocarpaceae dominated forest with the presence of some Castanopsis trees, in small groups on loamy soil, 17 July 2012, F. Hampe (GENT FH 12-158, duplicate: MFLU12-0648).Note Russula bellissima is a small species with a bright red pileus and pink colour on the stipe. This colour is distinctive and resembles North American R. mariae, Indian R. intervenosa and Asian R. bella. It is very unlikely that the distribution of any European or North American species is overlapping with the Thai species. However, little is known about the distributional ranges and the ecological niches of other Asian Russula species. Therefore discussing the morphological distinguishing characters between Asian species and R. bellissima is more relevant. Russula bellissima is not closely related to R. bella and it differs from this species by larger spores with a more prominent spore ornamentation, absence of hymenial cystidia on lamellae sides, and subterminally short, ellipsoid cells in the suprapellis arranged in unbranched chains of up to four7. The Thai species resembles and is closely related to the Indian R. intervenosa, but it has a more prominent spore ornamentation, hymenial cystidia (on lamellae sides) are absent, and hyphal terminations in the pileipellis are wider22.
Russula luteonana M. Pobkwamsuk & K. Wisitrassameewong sp. nov.
Mycobank: MB 840550Holotype: THAILAND, Amnat Charoen province, Hua Taphan district, Junction near Watbochaneng , dry dipterocarp forest, alt. 145 m, 15° 41′ 28″ N 104° 31′ 41″ E, 13 July 2016, Thitiya Boonpratuang, Rattaket Choeyklin, Prapapan Sawhasan, Maneerat Pobkwamsuk, Pattrachai Juthamas, Nattawut Wiriyathanawudhiwong, Patcharee Patangwesa (BBH41120).Etymology ‘Luteolus’ = yellow colour, ‘Nanus’ = small. Refer to pileus color and size of the species.Diagnosis Pileus medium-sized, dry, usually yellow, spores with subreticulate amyloid ornamentation and inamyloid suprahilar spot, hymenial cystidia on lamellae sides large, lamellae edges with combination of subulate, clavate and pyriform marginal cells.Pileus (Fig. 7) medium-sized, 28‒53 mm diam., plano-convex with depressed centre, infundibuliform when mature; margin striated and radially cracking in dry condition; cuticle dry, peeling to almost ½ of radius, smooth to minutely wrinkled, dull in dry condition, color very variable, some collections pale cream and with darker pale brownish-yellow centre, other yellow brownish and with darker orange-brown centre, sometimes also bright red-brown and with discolored centre, always with rusty-brown spots especially when near the centre. Lamellae: 3‒5 mm deep, moderately distant, intervenose, forking near the stipe, white to cream, edges even, concolorous. Stipe: 26‒40 × 6‒9 mm, cylindrical or narrowed at the base, surface dry, longitudinally wrinkled, white, turning brown when bruised. Context: 2‒4 mm in at the half pileus radius, soft, solid, becoming partially hollow when mature, white, unchanging when cut. Taste mild; odour rather strong, fishy. Spore print: not observed.Figure 7Basidiomata of Russula luteonana. (A) BBH41120 (Holotype). (B) BBH41121. (C) BBH41122. (D) BBH42510. Scale bar = 1 cm. Photos by Thitiya Boonpratuang.Full size imageSpores (Figs. 8, 9) (7.4‒)8.1‒8.6‒9(‒10.1) × (6.1‒)7.4‒7.5‒7.9(‒9.1) μm, subglobose to broadly ellipsoid, Q = (1.03‒)1.09‒1.15‒1.20(‒1.30), ornamentation of moderately distant, obtuse, (0.7‒)1.1‒1.3‒1.5(‒1.9) μm high spines, connected by abundant line connections [(0‒)3‒6(‒8) in in a 3 µm diam. circle], branched, forming an incomplete reticulum, crest irregularly wavy and occasionally fused [(0‒)1‒2(‒5) fusions in the circle], isolated elements rare; suprahilar spot inamyloid. Basidia: (29‒)34.5‒39.1‒44(‒51.5) × (10‒)12‒13.2‒14.5(‒16.5) μm, clavate, 4-spored, rarely 2-spored, basidiola subcylindrical to subclavate, (25.5‒)30‒35.4‒41(‒47) × (9‒)11‒12.2‒14 (‒16) μm. Hymenial cystidia on lamellae sides: usually protruding over other elements of hymenium, widely dispersed ( More

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Phyla Bdellovibrionota, Fusobacteriota, and Myxococcota were present in the green microbial mat but in negligible quantities in the brown mat. The unique phyla detected in the brown mat, but not in the green microbial mat, included Caldatribacteriota, Thermodesulfobacteriota, Dictyoglomota, Elusimicrobiota, Thermotogota, Candidatus Calescamantes, Fervidibacteria, Hydrothermae, GAL15 and TA06. The Candidatus Caldatribacterium (phyla Caldatribacteriota), earlier named OP9 was also detected in this work. Using single-cell and metagenome sequencing, data elucidated that Ca. Caldatribacterium conducts anaerobic sugar fermentation and exhibited diverse glycosyl hydrolases, including endoglucanase37.Cyanobacteria and Chloroflexota were the main identified phyla in the green microbial mat. Because the hot spring is almost stagnant, undisturbed, and the water surface temperature ( More