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Environmental Research
journal homepage: www.elsevier.com/locate/envres

Dynamic changes, cycling and downward fate of dissolved carbon and

nitrogen photosynthetically-derived from glaciers in upper Indus
river basin
Mohd Aadil Bhat a,b, Si-Liang Li b,c , Cong-Qiang Liu b,c , Nicola Senesi d, Giorgio S. Senesi e,
Davide Vione f,g , Daidu Fan a , Jie Yuan h , Mashura Shammi i , Khan M.G. Mostofa b,c,*
a
State Key Laboratory of Marine Geology, Tongji University, 1239 Siping Road, Shanghai, 200092, China
b
School of Earth System Science, Tianjin University, 92 Weijin Road, Tianjin, 300072, China
c
Tianjin Key Laboratory of Earth Critical Zone Science and Sustainable Development in Bohai Rim, Tianjin University, 92 Weijin Road, Tianjin, 300072, China
d
Dip.to di Scienze del Suolo, della Pianta e degli Alimenti, Università degli Studi di Bari "Aldo Moro", Via G. Amendola 165/A, 70126, BARI, Italy
e
CNR - Istituto per la Scienza e Tecnologia dei Plasmi (ISTP) - sede di Bari Via Amendola, 122/D - 70126 Bari, Italy
f
Università degli Studi di Torino, Dipartimento di Chimica, Via P. Giuria 5, 10125, Torino, Italy
g
Centro Interdipartimentale NatRisk, Via Leonardo da Vinci 44, 10095, Grugliasco, (TO), Italy
h
College of Resources and Environment, Xingtai University, Quanbei East Road 88, Qiaodong District, Xingtai City, Hebei Province, China
i
Department of Environmental Sciences, Jahangirnagar University, Dhaka, 1342, Bangladesh

A R T I C L E I N F O A B S T R A C T

Keywords: Glaciers play key roles in capturing, storing, and transforming global carbon and nitrogen, thereby contributing
Dissolved organic matter (DOM) markedly to their cycles. However, an integrated mechanistic approach is still lacking regarding glacier’s pri­
Dissolved organic nitrogen mary producers (PP), in terms of stable dissolved inorganic carbon isotope (δ13C-DIC) and its relationship with
Fluorescent DOM
dissolved carbon and nitrogen transformation d ynamic changes/cycling. Here, we sampled waters from glaciers,
Nutrients
δ13C-DIC
streams, tributaries, and the Indus River (IR) mainstream in the Upper IR Basin, Western Himalaya. Dissolved
Upper Indus river basin organic matter (DOM) appears to increase, on average, by ~2.5–23.4% with fluctuations when passing from
glaciers to streams-tributaries-IR mainstream (the upper and lower parts, respectively) continuum, implying that
DOM originates from glaciers PP and is subsequently degraded. The corresponding fluctuations are observed for
fluorescent DOM (FDOM), dissolved organic nitrogen (8.0–106.8%), NO−3 -N (− 13.5/+16.6%), NH+ 4 -N (− 8.8/
+13.0%), and NO−2 -N (70.7–217.5%). These variations are associated with overall DOM/FDOM transformations,
with the production of ending byproducts (e.g. CO2/DIC). The δ13C-DIC values fluctuated from glaciers (− 5.3 ±
2.5‰) to streams (− 4.4 ± 2.1‰), tributaries (− 4.3 ± 1.6‰), and IR mainstream (− 4.2 ± 1.3‰). The δ13C-DIC
data are consistent with C transformations that involve lighter CO2 emission into the atmosphere, whereas highly
depleted DIC/CO2 is the signature of DOM degradation after its fresh production from glaciers PP which orig­
inated by photosynthetic activities (e.g. uptake/sink of atmospheric CO2: − 8.4‰). Finally, glacier-fed meltwaters
would simultaneously contribute to the biogeochemical characteristics of downward margins and specific eco­
systems (lake/pond/groundwater/hot springs) via transformation dynamics/cycling of dissolved C and N with
high photo/microbial lability. Our results highlight the substantial contribution of western Himalayan glaciers-
derived DOM to the global C and N cycles.

1. Introduction which (77%) is in the dissolved organic carbon (DOC) form (Hood et al.,
2015), and are extremely vulnerable to climate change (Anesio et al.,
Globally, glaciers cover about 11% (734,400 km2) of the Earth’s land 2009; Zhou et al., 2019). Therefore, glaciers play a key role in the hy­
surface (Gardner et al., 2013), they store around 75% of the world’s drological cycle and substantially influence the global carbon (C) cycle
freshwater, contain about 6 Pg of organic carbon (OC), the majority of (Neckel et al., 2014; Hood et al., 2015). The Upper Indus River Basin

* Corresponding author. School of Earth System Science, Tianjin University, Tianjin, 300072, China.
E-mail address: [email protected] (K.M.G. Mostofa).

https://doi.org/10.1016/j.envres.2024.120117
Received 20 March 2024; Received in revised form 20 September 2024; Accepted 5 October 2024
M.A. Bhat et al.

(UIRB) is situated in the Himalaya and Karakoram (the H-K) mountain photosynthetically derived carbon in glaciers, based on the measure­
ranges, and includes almost half (47%) of the total Himalayan glacial ice ment of δ13C-DIC and carbon transformation dynamics in their down­
volume (Farinotti et al., 2019; Nie et al., 2021), and is considered the ward margins (streams, tributaries, and mainstream rivers). This
third pole of the Earth. It covers an area of 650,000 km2 (Himalaya) and knowledge gap also entails a lack of knowledge about the occurrence of
90,000 km2 (Karakoram), respectively, which extends across photosynthetically derived C in glaciers, and the role played by PP in
Afghanistan, Pakistan, China, India, Nepal, and Bhutan (Bolch et al., these ecosystems (Smith et al., 2016, 2017; Stibal et al., 2012; Foreman
2012; Bhat et al., 2021). Approximately, 869 million people living in the et al., 2013; Anesio et al., 2009; Hodson et al., 2010).
Indus, Tarim, Ganges, and Brahmaputra river basins rely on the water On the other hand, high atmospheric N depositions, particularly NH+ 4
and climate modulation controlled by the H-K (Nie et al., 2021). and NO−3 , in high-mountain Asia glaciers (Hattori et al., 2023; Gao et al.,
Glaciers are known to store dissolved organic matter (DOM) both on 2020; Huilin et al., 2008) are typically assimilated by glaciers PP and/or
their surface and subglacially, and then to transfer DOM after its gen­ partly removed/reduced via denitrification (Segawa et al., 2014; Tsu­
eration from photosynthetically-derived primary producers (PP, i.e., nogai et al., 2011; Murakami et al., 2022; Hattori et al., 2023). Such
cyanobacteria, algae, and diatoms) (Musilova et al., 2017; Pautler et al., glaciers PP primarily release the proteinaceous substances, autochtho­
2012; Smith et al., 2017; Hood et al., 2009; Stubbins et al., 2012; nous humic-like substances, and pigments (Smith et al., 2017; Zhou
Spencer et al., 2014; Lawson et al., 2014a). Notably, the DOC stored in et al., 2019; Stubbins et al., 2012; Spencer et al., 2014; Barker et al.,
glaciers amounts to approximately 4.48 ± 2.79 Pg C, which represents 2013; Feng et al., 2021), which are generally composed of dissolved
about 75% of the total OC stored in glaciers and ice sheets (Hood et al., organic nitrogen (DON) (Zhou et al., 2019; Gao et al., 2020; Stedmon
2015). Glacier DOM has been widely studied in Antarctica and the Arctic et al., 2007a). Such DON-bound DOM components are rapidly degraded
(Hood et al., 2015; Pautler et al., 2012; Smith et al., 2017; Barker et al., into NH+ 4 and other byproducts, either photochemically and/or micro­
2013), Canada and the USA (Stubbins et al., 2012), Greenland ice sheet bially (Stedmon and Markager, 2005; Stedmon et al., 2007b; Smith
(Hood et al., 2015; Musilova et al., 2017; Lawson et al., 2014a, 2014b; et al., 2017), which could partly be exported into glaciers-fed meltwa­
Bhatia et al., 2010), Tibetan Plateau (Zhou et al., 2019; Spencer et al., ters or used by glaciers PP. Moreover, total nitrogen (TN) has been
2014; Feng et al., 2021), Alaska (Hood et al., 2015; Stubbins et al., 2012) studied in isolated DOM molecules from glaciers (Feng et al., 2021),
and the European Alpine glaciers (Singer et al., 2012). The chemical together with the cycling of either N-nutrients (Smith et al., 2016) or
composition of DOM in these areas has been characterized at the mo­ DON in glaciers and glaciers-fed meltwaters in streams, and rivers (Zhou
lecular level by FT-ICRMS (Zhou et al., 2019; Stubbins et al., 2012; et al., 2019). However, a gap still remains regarding the dynamic
Spencer et al., 2014; Feng et al., 2021), and it has been shown to be changes (i.e., decrease and/or increase) of TN and related N-nutrients in
primarily composed of proteinaceous substances, autochthonous glaciers and downward streams, tributaries, and mainstream river wa­
humic-like substances, and pigments (Zhou et al., 2019; Smith et al., ters in the large UIRB basin. It is well known that DOM and particularly
2017; Stubbins et al., 2012; Spencer et al., 2014; Barker et al., 2013; fluorescent DOM (FDOM) components are consistently altered in gla­
Feng et al., 2021). In particular, a number of earlier studies have been ciers and their meltwaters, in the stream channels, and along their
conducted in the UIRB which mostly include: (i) hydrological studies, margins (Smith et al., 2017; Barker et al., 2013). Moreover, limited
with measurements of moisture sources, hydrochemical processes, and knowledge is available on DOM and FDOM transformation dynamics, i.
chemical weathering processes (Bhat et al., 2021, 2023; Lone et al., e., on the production of DOM and FDOM from glaciers PP and its sub­
2020, 2021); (ii) glaciological studies, with monitoring glacier mass sequent degradation, as well as the relationships with nutrient transport
balance and retreat rates, highlighting the critical role of glaciers in and dynamics, particularly with regard to N-nutrients.
sustaining river flow (Bolch et al., 2012, Lone et al., 2023); (iii) climatic Based on the above considerations, the primary objectives of this
studies, to assess the impacts of global warming on glacial melt and study are to: (a) ascertain the distribution and potential sources/sinks of
precipitation patterns, revealing significant changes that affect water DOM release from photosynthetically-derived PP (e.g. phytoplankton)
resources (Lutz et al., 2014); and (iv) environmental studies, with in glaciers, and its subsequent transformation upon transport from gla­
measurement of heavy metals, to ascertain the impacts of pollution and ciers to streams, tributaries, and the Indus River (IR) mainstream waters,
human activities on a specific river (e.g. Hunza River and its tributaries) in order to better understand dissolved C and N cycling by compre­
ecosystem (Ali et al., 2015). However, DOM dynamic changes/cycling hensively measuring δ13C-DIC for the first time and N-nutrients in the
together with its fluorescence characterization in the UIRB remain UIRB; (b) characterize FDOM components using the EEM-PARAFAC
elusive. model, and discuss their alterations in glaciers, streams, tributaries,
The stable carbon isotope (δ13C) of DIC has been used as an efficient and the IR mainstream waters; and (c) Evaluate the influence of glaciers-
tracer to quantify DIC sources/sinks and transformation processes in fed meltwaters on the biogeochemical characteristics of specific eco­
surface waters (Marwick et al., 2015; Yang et al., 1996; Yi et al., 2021). systems (a saline lake, a pond, groundwaters, and hot springs) in the
Furthermore, DOM components exhibit rapid degradation by hetero­ UIRB, by measuring DOM, FDOM, δ13C-DIC, and nutrients. Further­
trophs (Smith et al., 2017), as well as upon sunlight exposure (Yi et al., more, the results of this study provide a conceptual model that depicts
2021; Yang et al., 2021), which causes an enrichment in δ13C (Pautler the production processes of photosynthetically-derived C in glaciers and
et al., 2012; Smith et al., 2017; Hood et al., 2009), and δ15N (Lawson roughly estimates the C and N cycling during the transformation of
et al., 2014b). Degradation processes release CO2, dissolved inorganic DOM/FDOM, δ13C-DIC, and nutrients under photoinduced and/or bio­
carbon (i.e., DIC, which includes CO2 + H2CO3 + HCO−3 + CO2− 3 ), nu­ logical respiration in glaciers and along their margins.
trients, and low molecular weight DOM (Stedmon et al., 2007b; Bhatia
et al., 2010; Stubbins et al., 2010; Singer et al., 2012), which involve 2. Materials and methods
CO2 exchange at the water-air interface with subsequent impact on C
isotopic fractionation. In principle, the uptake of lighter/enriched 2.1. Site description
δ13C-DIC/CO2 by aquatic photosynthetic PP (e.g. phytoplankton) that
are part of the particulate organic carbon (POC) could generate light­ The Indus River (IR) originates from Mount Kailash near Mansarovar
er/enriched δ13C-POC, whereas lighter δ13C-POC could produce lighter Lake in Tibet (China) and is one of the largest drainage basins on the
δ13C-DOC that undergoes substantial degradation (Smith et al., 2017; Yi Earth, featuring an area of 1,165,500 km2 (Bhat et al., 2021). It com­
et al., 2021), with subsequent production of lighter/enriched δ13C-DIC prises the high-altitude Himalayan valleys and the Karakoram mountain
and corresponding enriched δ13C-DOC (Yi et al., 2021; Pautler et al., ranges in India, crosses Pakistan, and drains into the Arabian Sea. The
2012; Smith et al., 2017; Hood et al., 2009). However, currently there is study area is situated in northwest (NW) India and is referred to as
no integrated understanding of sinks/sources in terms of Trans-Himalayas, covering an area from 75◦ 30′32′′ - 78◦ 56′18″E and
M.A. Bhat et al.

33◦ 04′18′′ - 34◦ 60′38″N (Fig. 1). The UIRB has a total length of ~429 km, ensure sample homogeneity. Glacier ice samples were allowed to melt at
and a watershed area of approximately 53,567 Km2 (Bhat et al., 2021, room temperature and then stored in 30 mL HDPE narrow-headed,
2023). Further site descriptions are provided in the Supporting Infor­ tightly capped sampling bottles. The water samples from streams, trib­
mation (SI: subsection 2.1). utaries, and the IR mainstream were collected from the flowing areas
near the center of each sampling site. They were collected from the
2.2. Sample collection and preparation uppermost surface (approximately <0.5 m), mostly from the middle of
the IR mainstream using a long rope connected to a polycarbonate
Water samples were collected from glaciers and their downward bucket (over the bridges). In some cases, in the absence of bridges, we
margins (streams, tributaries, and the IR mainstream, as well as glaciers- used small boats to reach the middle river or threw a polycarbonate
fed specific ecosystems (Pangong Tso saline lake, pond, groundwaters, bucket into the middle connected with a rope, while staying along the
and hot springs) of the UIRB between July and August 2019 (Fig. 1; sides/bank of the river (Fig. 1; Table S1). Water samples from saline lake
Table S1). The glacier ice sampling was performed in snouts of the and pond were collected along the sides by throwing a polycarbonate
Khardungla, Drungdung, and Changla groups of glaciers in the Indus bucket into the middle, connected with a rope. Water samples from hot
Basin, and meltwater was sampled at the base of the glacier pack to springs were directly collected, at approximately 3–4 cm depth.

Fig. 1. Topography of locations of sampling sites in the Upper Indus River Basin (UIRB), Ladakh.
M.A. Bhat et al.

Groundwater samples were collected after 10–15 min of pumping from ratio mass spectrometer (IRMS) (Thermo Fisher, 0.3‰, analytical
wells located close to the rivers/streams, near the towns of Drass, Kargil, error)2 and expressed in ‰ of the international V-PDB standard. The
and Leh. A portable hand-held, multi-parameter kit (Hanna, In­ remaining information regarding site description, fluorescence spec­
struments, USA), which was calibrated before sampling, was used to troscopy and PARAFAC modeling, dissolved organic nitrogen (DON)
measure the pH, temperature, and electrical conductivity of the samples calculation, CO2 and total nitrogen (TN) flux calculations, statistical
in field conditions. A portable global positioning system (GPS, GARMIN, data analysis, and GIS, as well as the changes in the biogeochemical
eTrex30) was used for geo-reference of each sampling location. Then, properties of glaciers-fed specific ecosystems, are outlined in the SI
water samples were filtered through a 0.45-μm glass-fiber filter (GF/F, (subsections 2.1, 2.4–2.7.1, and 3.2.).
Whatman, Inc.) and stored in acid-washed, high-density polyethylene
bottles, which were kept refrigerated at 4 ◦ C until further analysis. 3. Results and discussion

2.3. Chemical and isotopic analyses 3.1. Transportation and transformation dynamics from glaciers to
streams, tributaries, and the IR mainstream
The DOC concentration was measured by an Aurora 1030 total
organic carbon (TOC) analyzer (OI Analytical Aurora, Model 1030W + 3.1.1. DOC
1088, USA) after acidification with HCl to remove all inorganic carbon. The DOC content varies from 1.1 to 2.6 mg/L (average: 1.7 ± 0.32
At least three injections were performed for each DOC analysis and mg/L) in glaciers, streams, tributaries, and the IR mainstream water
potassium hydrogen phthalate was used as the DOC standard solution. samples (Table S1). On average, a slight increase (7.9%) is detected
The concentrations of nutrients (NO¡ 3 -N, NH4 -N, NO2 -N, PO4–P, and
+ ¡
when passing from glaciers to streams, then a decrease in tributaries,
dissolved Si) were measured by a Skarlar San continuous flow online and again an increase (23.4%) in the IR mainstream (Fig. 2, Fig. S1A,
analyzer (Skarlar Company, ++ Breda, Netherlands) (Yang et al., 2020; Table S2). Such DOC variations are statistically significant in a two-
Yang et al., 2021). sample z-test only for glaciers to the IR mainstream (the upper and
For the determination of δ13C-DIC, water samples were first injected lower parts) and for tributaries to the IR mainstream (the upper and
by a syringe into a vacuum bottle containing 85% phosphoric acid and a lower parts) (Table S3). These results suggest an additional input to
bar magnet. Then, the produced CO2 was extracted and cryogenically DOM originated from the autotrophic activity of microbial communities
purified in a vacuum system (Zhong et al., 2018). After CO2 purification, in glacial-fed meltwaters (Pautler et al., 2012; Antony et al., 2012;
δ13C-DIC was measured using a Delta-V + Gas Bench II stable isotope Hodson et al., 2010; Stibal et al., 2012), and/or sunlight-induced

Fig. 2. Box plots representing the minimum and maximum values of dissolved organic carbon (DOC) concentration in waters collected from glaciers, streams,
tributaries, the Indus River (IR) mainstream (upper part), the IR mainstream (lower part), hot springs, saline lake, pond, and groundwaters in the Upper IR Basin
(UIRB) during the summer season. The solid line inside the boxes represents the median and error bars represent the standard deviation (SD) of the mean.
M.A. Bhat et al.

respirations (Yang et al., 2021). This agrees with earlier studies into This finding also agrees with earlier studies (Cory et al., 2014; Smith
downstream water transport (Smith et al., 2016, 2017; Stibal et al., et al., 2017; Stedmon et al., 2007b; Zhou et al., 2019; Panwar et al.,
2012; Foreman et al., 2013; Anesio et al., 2009). Importantly, the gla­ 2017). Furthermore, the presence of barren catchments, of poorly
ciers DOC concentrations (average: 1.53 ± 0.14 mg/L) of the UIRB were developed soils and vegetation, and the lower amount of rainfall in this
relatively higher compared to those of other high-mountain glaciers, area (Lone et al., 2021) are not expected to contribute to terrestrial DOC
such as 0.28 ± 0.20–1.17 ± .34 mg/L in Dongkemadi/Geladaindon­ input into rivers. Additionally, the lithology of the surrounding UIRB
g/Laohugou/Baishui glaciers, Tibetan Plateau (Li et al., 2018); consists largely of carbonates and other organic-poor sedimentary rocks
0.52–0.35 ± 0.09 in Ürümqi glacier, Tienshan; 0.81 mg/L in East (Bhat et al., 2023; Lone et al., 2020), which also do not significantly
Rongbuk glacier, Mt. Everest, Himalayas, and 0.24 mg/L in Joyce contribute DOC to river waters.
Glacier, Garwood Valley, Antarctica (reviewed in Gao et al., 2020);
0.12–0.43 mg/L in John Evans glacier, Canada (Barker et al., 2006); 3.1.2. Nutrients
0.25–1.18 mg/L in Austre Brøggerbreen glacier, Svalbard (Zhu et al., The spatial distributions of physico-chemical properties and all nu­
2016); and 0.21 ± 0.04 mg/L in snowpit of Muz Taw glacier in Central trients are shown in Fig. S2. Glaciers typically feature relatively high
Asia which, however, showed higher values in the surface aged snow NO−3 -N and DSi, as well as relatively low NH+ 4 -N, NO2 -N, and PO4-P
−

(1.12 ± 1.66 mg/L) (Gao et al., 2020), comparable to values found by us (Tables S1 and S4, Fig. 4). On average, TN increased from glaciers to
in UIRB glaciers. streams, tributaries, the IR mainstream (the upper part), and the IR
Furthermore, the DOC content in the right-side streams of the gla­ mainstream (the lower part) by, respectively, 12.7%, 6.7%, 14.5%, and
ciers is slightly higher, by approximately 9.5%, than in the left-side 8.8% (Table S2). These results might be associated with diverse sce­
streams, which could be ascribed to in-situ production of DOM from narios involving DON and N-nutrients, which include basin runoff and
PP via intense photo-microbial respirations relatively far from glaciers, atmospheric N2 fixation (Stedmon et al., 2007b; Zhang et al., 2021;
due to sunrise impinging right-side stream waters and to relatively Hattori et al., 2023; Gao et al., 2020; Huilin et al., 2008; Telling et al.,
higher water temperature (WT, 14.4–19.2 ◦ C, average: 16.1 ± 2.2 ◦ C), 2011). The TN content is accounted for by 62.0–72.3% NO−3 -N,
compared to the case of left-side streams (WT of 10.7–24.8 ◦ C, average: 18.0–26.6% DON, 11.3–16.7% NH+ 4 -N, and 0.9–2.6% NO2 -N, with some
−

14.7 ± 4.9 ◦ C; Table S4). Differently, the average DOC content was variations between glaciers and streams, tributaries, and IR mainstream.
similar (1.57 mg/L) for right-side and left-side tributaries. The upper These results would suggest that DON and NO−3 -N play an important role
part of the IR mainstream shows DOC values 4.6% higher than the lower in N-nutrient cycling. Compared to TN, the single N components (NO−3 ,
part, which might mostly be due to the higher WT (14.5–32.0 ◦ C, DON, NH+ 4 , and NO2 ) show much larger fluctuations, which would
−

average: 23.5 ± 5.1 C, IR-sites 1–8 vs 14.1–18.5 ◦ C, average: 15.4 ± suggest the occurrence of nitrification-denitrification processes during
○

1.3 C), possibly occurring intense photo-microbial respirations of downstream water transport, in agreement with earlier studies (Segawa
○

photosythetically-derived PP in glaciers-fed meltwaters under higher et al., 2014; Hattori et al., 2023; Van Kessel et al., 2015; Ward et al.,
WT conditions (Smith et al., 2017; Yang et al., 2021). This agrees with 2009). Remarkably, a substantial increase in DON contents (primarily
relatively depleted δ13C-DIC values (average: − 4.4 ± 0.8‰) in the upper associated with the molecular configuration of DOM) in the upper part
part of the IR mainstream compared to the lower part (average: − 4.0 ± of the IR mainstream compared to tributaries and the lower part of the
1.5‰), which is the key signature of newly-derived DOM export from IR mainstream (Table S2) is a key signature of increasing DOM contents
glaciers PP, which also agrees with earlier studies (Table S1; Cory et al., in the upper part, as discussed in ealier section. These results are further
2014; Smith et al., 2017; Yang et al., 2021). Such high confirmed by the positive correlation of DOC with TN, DON, NH+ 4 -N, and
temperature-dependent DOM export is further supported by the Pearson NO−2 -N (Table S5), as well as by PCA where PC1 includes high positive
correlation between DOCand WT (Table S5), as well as by Principal loadings for WT (T◦ C, 0.756), DOC (0.700), TN (0.704), NO−3 -N (0.496),
Component Analysis (PCA) in which PC1 explains 23.6% of the total NH+ 4 -N (0.524), and DON (0.381) (Table S2). These results suggest the
variance and is characterized by high positive loadings for WT (T◦ C, occurrence of changes in DON-bound DOM induced by warming, as well
0.756) and DOC(0.700; Table 1; Fig. 3A), indicating a potential relation as the cycling of nutrients. PC1 results are further emphasized by PC2
between warmer waters and high DOM levels. Differently, relatively that accounts for 15.6% of the variance, with the strongest positive
enriched δ 13C-DIC values in the lower part of the IR mainstream are loadings for DON (0.724) and TN (0.539), and negative loadings for
presubambly due to sequential degradative loss of lighter C (Table S1). PO3−4 -P (− 0.603), NH4 -N (0.403) and NO2 -N (0.413) (Table 1), which
+ −

implies a strong relation between organic and inorganic forms of ni­

trogen and phosphorus.
Table 1 Similarly, DSi substantially increases, on average, by approximately
Loadings of 12 environmental parameters on five significant principal compo­ 130%, 141%, 506%, and 226% in streams, tributaries, and the IR
nent variables and total variance explained, for samples collected from glaciers, mainstream (the upper and lower parts, respectively) with respect to
streams, tributaries and the Indus River (IR) mainstream in the upper IR Basin glaciers (Table S2). These results might most likely be ascribed to Si
(UIRB), western Himalaya. Bold values indicate strong and moderate release in the form of, e.g., H4SiO4 from silicate minerals by chemical
associations.
weathering (Bhat et al., 2023; Hatton et al., 2019). On average, PO4-P
Variables PC1 PC2 PC3 PC4 PC5 increases by 27.5%, 108.0%, 118.8%, and 6.3% in streams, tributaries,
T C
◦
0.756 − 0.353 − 0.224 − 0.033 − 0.007 and the IR mainstream (the upper and lower parts, respectively),
pH 0.050 − 0.316 0.086 0.776 − 0.048 compared to glaciers (Table S2). Similar ranges of PO4-P values are
EC 0.081 − 0.261 0.299 − 0.142 0.736 typically detected in glacier-fed estuarine waters (Lund et al., 2018).
DOC 0.700 0.040 − 0.226 0.025 − 0.400
These results suggest that the increase and subsequent decrease of PO4-P
δ13C-DIC − 0.003 − 0.218 0.552 0.546 − 0.178
TN 0.704 0.539 0.408 − 0.032 − 0.003 would be most likely associated with the photochemical and microbial
DON 0.381 0.724 − 0.119 0.172 0.213 degradative release from DOM and with photosynthetic activity (Lund
NO3-N 0.496 0.127 0.713 − 0.188 − 0.185 et al., 2018; Liu et al., 2017; Sosa et al., 2019; Repeta et al., 2016).
NH+4 -N 0.524 − 0.403 − 0.311 0.184 0.084
NO2-N 0.423 − 0.413 − 0.091 − 0.354 − 0.309
PO4-P 0.116 − 0.603 0.376 − 0.257 0.135
3.1.3. δ13C− DIC characteristics and its potential sources
DSi 0.643 0.021 − 0.214 0.179 0.486 The δ13C-DIC values vary from − 9.5 to − 3.1‰ (average, − 5.3 ±
Eigenvalue 2.837 1.869 1.484 1.246 1.172 1.7‰) in glaciers and, on average, they are slightly enriched in streams
Variability (%) 23.641 15.575 12.367 10.381 9.767 (− 4.4 ± 2.1‰) and tributaries (− 4.3 ± 1.6‰), and then depleted and
Cumulative % 23.641 39.216 51.584 61.964 71.732
enriched again in, respectively, the upper part and the lower part of the
M.A. Bhat et al.

Fig. 3. Biplots of principal component analysis (PCA) illustrating the loadings of relevant experimental parameters between significant principal components (PC1
and PC2: A) for samples collected from glaciers, streams, tributaries, and the mainstream of the Indus River (IR); and significant principal components (PC1 and PC2:
B) for samples collected from glaciers, hot springs, saline lake and groundwater in the upper IR Basin, western Himalayas.

IR mainstream (− 4.4 ± 1.0‰ and − 4.2 ± 1.5‰, respectively) freshly produced upon uptake of atmospheric CO2 (− 8.4‰ in 2015;
(Tables S1 and S4; Figs. S1B, 4A, B). Such variations are not statistically Graven et al., 2020) in conditions of alkaline pH (8.2–9.1). This would
significant when performing a two-sample z-test (Table S3). Impor­ be caused by the high solar intensity in the glacier site compared to the
tantly, the Khardunga top glacier shows the most depleted δ13C-DIC rest of India (see details in site description, Text SI-2.1), ambient air
(− 9.5‰), which is followed by a gradual increase in the downward di­ temperature (AT) below zero, and low WT (8–11 ◦ C).
rection, i.e., in the glacier mid-point and end/last point (melt waters; The production of OM and the corresponding DOM production in
respectively, − 4.7‰ and − 3.1‰). These findings suggest that glaciers are widely reported by earlier studies (Smith et al., 2016, 2017;
photosynthetically-derived PP/organic matter (OM) in the top glacier is Singer et al., 2012; Stibal et al., 2012; Foreman et al., 2013; Hodson
M.A. Bhat et al.

Fig. 4. Concentrations of various nutrients, i.e., total nitrogen (TN), dissolved organic nitrogen (DON), NO3-N, NH+ 4 -N, NO2-N, PO4-P, and dissolved Si (DSi) in
waters collected from glaciers, streams, tributaries, and the IR mainstream (upper and lower parts: A-B) and various ecosystems, i.e., hot springs, saline lake, pond
and groundwaters (C–D) in the Upper Indus River Basin (UIRB) during the summer season. Error bars represent the standard deviation of the average values of
each parameter.

et al., 2010). The subsequent degradation of DOM by photo/microbial highlight the transformation of δ13C-DIC and NO−3 , with the corre­
respiration generates depleted δ13C-DIC including depleted CO2 sponding warming-induced alterations of DOM and NH+ 4 -N. PC3 results
(− 9.5‰) in glaciers (Stedmon et al., 2007b; Bhatia et al., 2010; Stubbins are further emphasized by PC4, which accounts for 10.4% of the vari­
et al., 2010; Singer et al., 2012; Bade et al., 2004). In particular, the ance and shows significant positive loadings for δ13C-DIC (0.546) and
highly depleted δ13C-DIC (− 9.5‰) in the Khardunga top glacier implies pH (0.776), as well as negative loadings of NO−2 -N (Table 1), suggesting
that lighter CO2/DIC is freshly originated from glacier DOM and has not that δ13C-DIC transformation might be influenced by pH and NO−2 -N.
yet been emitted into the atmosphere by hydrological processes due to Overall, the δ13C-DIC values measured in UIRB are similar to those in
its occurrence in the top glacier (Bade et al., 2004; Lammers et al., 2017; some relevant world rivers (e.g. Yangtze, Yellow, Ganga, and Indus),
Bhat et al., 2021; Yi et al., 2021). This result is further supported by the which is presumably due to similar sources of photosynthetically-
fact that ice sheet interiors would function as sinks for atmospheric CO2, derived DOM, but dissimilar to those of other important rivers (e.g.
whereas the ice sheet edges and small glaciers would act as sources Amazon, Mekong, Ottawa, Rhine, and Mascouche) (Fig. 5C). The dif­
(Stibal et al., 2012). Differently, the enrichment of δ13C-DIC in down­ ferences might be ascribed to diverse sources of DIC, e.g., terrestrial,
ward glaciers-fed meltwater would arise from lighter CO2 emission into anthropogenic, or photosynthetically-derived DOM (Cai et al., 2008; Li
the atmosphere by hydrological processes/evaporation (Morana et al., et al., 2010; Singh et al., 2005; Stedmon et al., 2007b; Stubbins et al.,
2015; Yi et al., 2021; Bade et al., 2004), and not by photosynthetic 2010; Singer et al., 2012; Bade et al., 2004). Based on their isotopic
uptake due to water movement. These results confirm those of our signature, the UIRB samples appear unaffected by anthropogenic sour­
earlier study, showing that the hydrological processes alter the isotopic ces and regulated only by glaciers-fed meltwaters (Stubbins et al., 2010;
signatures of δ13C-DIC and δ18O when passing from the Khardungla top Singer et al., 2012; Bade et al., 2004). Therefore, the primary source of
to its two downward sites (Bhat et al., 2021). CO2/DIC in UIRB is the DOM/FDOM that is produced by transformation
Similarly, highly depleted and highly enriched δ13C-DIC values are (upon photoinduced and microbial respiration) of
often detected in individual streams, tributaries, and the IR mainstream photosynthetically-derived DOM in glaciers, a transformation that
(Fig. 5A and B), which is possibly due to the high production of lighter would occurs when passing from glaciers to streams, tributaries, and the
CO2/DIC from OM and/or emission of lighter CO2 to the atmosphere by IR mainstream (Stibal et al., 2012; Li et al., 2010; Singh et al., 2005).
hydrological processes/evaporation, depending on the riverine site. This Photoinduced and microbial processes together with evaporation/hy­
interpretation is further supported by the strong correlation (r2 = 0.44, drological processes appear thus to be the key regulators in enriching
p < 0.01) between DOCand δ13C-DIC (Table S5), which is also consistent δ13C-DIC during water transport from glaciers to their downward mar­
with the positive correlation between OM bioavailability and in-stream gins, with continuous emission of lighter CO2 to the atmosphere, thereby
CO2 concentrations, as previously found (Singer et al., 2012). Moreover, constantly contributing to the global C cycle.
the average enrichment of δ13C-DIC values in the lower part of the IR
mainstream compared to its upper part (Fig. 5B; Table S4) would arise 3.1.4. Characterization of fluorescent DOM (FDOM)
from emission of lighter CO2/DIC to the atmosphere, whilst a substantial The EEM-PARAFAC model identified two to six fluorescent compo­
δ13C-DIC depletion in the middle part of the IR mainstream (Fig. 5A) nents (Fig. 6, Fig. S5) in the examined individual samples. The corre­
would arise from an extended CO2/DIC production from DOM degra­ sponding Ex/Em peak positions are summarized in Table S6 and their
dation, after the confluence mixing of the major tributary, i.e. the peak intensities are provided in Fig. S6. Components one (C1) and two
Zanskar river with the IR mainstream, on a gravel-mixed riverbed. Such (C2) are universally assigned to autochthonous humic acid-like sub­
changes in the δ13C-DIC values from glaciers to streams, tributaries, and stances (AHLS), each of which features two fluorescence peaks, i.e., A-
the IR mainstream are partially explained by PCA where PC3 explains type and C-type, as well as C-type and M-type peaks, respectively. These
12.4% of the variance, with significant positive loadings for δ13C-DIC two components (C1 and C2) are primarily derived from
(0.552) and NO−3 -N (0.713), as well as negative loadings for DOC photosynthetically-derived PP/OM (e.g. phytoplankton) in glaciers-fed
(− 0.226), NH+ 4 -N (− 0.311), and WT (− 0.224) (Table 1). These data meltwaters, which agrees with many field and experimental
M.A. Bhat et al.

Fig. 5. (A) Variation of the values of stable isotope of dissolved inorganic carbon (δ13C-DIC) values in the Indus River (IR) mainstream in the Upper IR Basin (UIRB).
(B) Box plots representing the minimum and maximum δ13C-DIC values in waters collected from glaciers, streams, tributaries, the IR mainstream, hot springs, saline
lake, pond, and groundwater in UIRB during the summer season. Error bars represent the standard deviation (SD) of the mean. (C) Comparison of the minimum and
maximum δ13C-DIC values of all samples in UIRB with the 15 largest world rivers. The solid line inside the box in (b) and (c) represents the median. The δ13C-DIC
literature data sources are: atmosphere (Doctor et al., 2008; Dubois et al., 2010); Amazon River (Marwick et al., 2015); Yangtze River (Shan et al., 2021); Brah­
maputra River (Singh et al., 2005); Mississippi River (Dubois et al., 2010); Mekong River (Li et al., 2019); Ganga River (Singh et al., 2005; Chakrapani and Veizer,
2005); St Lawrence River (Hélie et al., 2002); Pearl River (Shan et al., 2021); Heilongjiang River (Shan et al., 2021); Yellow River (Shan et al., 2021); Indus River
(Karim and Veizer, 2000); Ottawa River (Hélie et al., 2002); Rhine River (Flintrop et al., 1996); Mascouche river (Hélie et al., 2002).

observations (Zhou et al., 2019; Yang et al., 2021; Zhang et al., 2009; sedimentary rocks (Bhat et al., 2023; Lone et al., 2020) and there was
Shammi et al., 2017a), as well as OpenFluor database (Stedmon et al., no rain before and during the sampling period. Component three (C3) is
2007a, 2007b; Chen et al., 2018; Gao and Guéguen, 2017; Wünsch et al., assigned to a combined form of C-type and M-type AHLS (Nye et al.,
2018; Dainard et al., 2019; DeFrancesco and Guéguen, 2021; Imbeau 2020; Shammi et al., 2017b; Yang et al., 2021), each of which features
et al., 2021; Kurek et al., 2022). Notably, they are not mixing up with two fluorescence peaks and is present only in waters from hot springs
allochthonous/terrestrial sources because the lithology of the sur­ (discussed in SI, subsection 3.2.4), which also agrees with OpenFluor
rounding UIRB largely consists of carbonates and other organic-poor database (Chen et al., 2018; Gonçalves-Araujo et al., 2015; Wünsch
M.A. Bhat et al.

Fig. 6. Fluorescent components identified by the PARAFAC model in water samples collected in the Upper Indus River Basin UIRB), where component C1: C-type
autochthonous humic acid-like (C-type AHLS); C3: a combined form of C-type and M-type AHLS; C4: tryptophan-like substances (TLS); C5: newly-released protein-
like substances (PLS); C6: tyrosine-like substance (TYLS) and/or phenylalanine-like substance (PALS); and C7: a degraded byproduct of methanol. Note that C3 has
been detected only in hot spring waters (SI, subsection 3.2.4 and Fig. S5). Excitation and emission loadings produced from the respective PARAFAC models for the
respective individual samples are also presented in Fig. S5.

et al., 2017; Gao and Guéguen, 2017; Maurischat et al., 2022). and Scott, 2005; Ebinghaus and Wilken, 1993).
Component four (C4) is attributed to tryptophan-like substances (TLS) Five fluorescent components are identified in glaciers, which feature
and features two peaks (T and TUV) (Zhou et al., 2019; Coble, 1996), EEM spectra (Fig. 6) and corresponding peaks (Table S6) that suggest a
which also agrees with OpenFluor database (Stedmon et al., 2011; phytoplankton source (Stubbins et al., 2012; Zhou et al., 2019; Yang
Jørgensen et al., 2011; Wünsch et al., 2017; Kida et al., 2019; DeFran­ et al., 2021; Yi et al., 2021; Yamashita and Tanoue, 2003), and that
cesco and Guéguen, 2021; Lu et al., 2022; Maurischat et al., 2022; differ from soil humic substances (HS), including humic acids, fulvic
Drozdova et al., 2022). Component five (C5) is ascribed to acids, and PLS PLS (Gao et al., 2018; Mohinuzzaman et al., 2020). This
newly-released protein-like substances (PLS) and consists of two peaks interpretation is supported by the depleted radiocarbon of glacier DOM,
(Zhou et al., 2019; Shammi et al., 2017b; Yang et al., 2021), which also which suggests a microbial and/or anthropogenic source with no
agrees with OpenFluor database (Walker et al., 2009; Kowalczuk et al., terrestrial contribution (Stubbins et al., 2012; Foreman et al., 2013;
2013; Brogi et al., 2019; DeFrancesco and Guéguen, 2021; Maurischat Barker et al., 2013; Smith et al., 2017). The glacier component C2 would
et al., 2022). Component six (C6) is assigned to tyrosine-like substances be a newly derived M-type AHLS, which exhibits three fluorescence
(TYLS) and/or phenylalanine-like substancess (PALS) and features two peaks, i.e., peak M, 305/419, 265/419 and peak A, 235/419 nm, which
peaks (Yamashita and Tanoue, 2003; Yang et al., 2021), which also would be freshly originated from phytoplankton OM by photoinduced
agrees with OpenFluor database (Jørgensen et al., 2011; Painter et al., and/or microbial respiration, at a WT of 8.0–10.5 ◦ C and a pH of
2018; D’Andrilli et al., 2017). Component seven (C7) features two peaks 8.3–9.1. This interpretation is further supported by the slow production
(T: 245–250/284~290 nm and TUV: 220/284–290 nm; Table S6), and is of AHLS with PLSsignatures, as in the case of intense TYLS fluorescence
considered to be a degraded byproduct of various FDOM types, typical of from filamentous cyanobacteria and diatoms, detected during incuba­
either methanol derivatives as compared to their standards in aqueous tion experiments experiments (Smith et al., 2017). Three similar peaks
solution (Table S6; Fig. S5; Parks et al., 2013), or hydrolysis products of have also been detected in resuspensions of a lake algal biomass in
methyl derivatives present in surface waters (e.g. CH3I, CH3Hg) (Celo Milli-Q water under dark incubation (Mostofa et al., 2013). Apart from
M.A. Bhat et al.

component 7, the other three components (TLS, newly-released PLS, and nutrients between glaciers and individual specific ecosystems are evi­
TYLS) can be related to proteinaceous materials that are key signatures denced by the results of PCA, where PC1 accounts for 38.0% of the
of autochthonous DOM sources (Smith et al., 2016, 2017; Stibal et al., variance and is characterized by high positive loadings of electric con­
2012; Foreman et al., 2013; Anesio et al., 2009) and would be derived ductivity (EC, 0.922), DOC (0.919), DON (0.921), δ13C-DIC (0.892), and
from the autotrophic activity of microbial communities (Pautler et al., pH (0.799), along with negative loadings of NO−3 -N and PO3−4 -P (Table 2;
2012; Hodson et al., 2010; Stibal et al., 2012). Fig. 3B). These results suggest that a significant increase and/or trans­
The Indus Valley stream (IVS) samples show six fluorescent compo­ formation of organic C plus N in DOM, and of δ13C-DIC together with EC
nents, of which C1 (C-type AHLS) and C2 (M-type AHLS) can be and pH could arise together with consumption of nutrients (e.g. NO−3 -N
considered as derivatives of glaciers freshly-derived C2 (M-type AHLS), and PO3−4 -P). Such changes in C and N are further supported by the PC2
at a WT of 11.8–22.7 ◦ C and pH of 8.3–9.2. The other four components and PC3 results, which are substantially influenced by temperature and
are similar to those of glaciers (Fig. 6, Table S6). The stepwise produc­ by the corresponding increase of some nutrients (Table 2). Notably, PC2
tion of C-type and M-type AHLS from combined forms has been hy­ explains 25.0% of the variance and is characterized by high positive
pothesized by Yang et al. (2021), and Smith et al. (2017) who showed loadings of WT (0.917), DSi (0.976), NH+ 4 -N (0.683), and NO2 -N
−

the occurrence of an increasing humic-like fraction from proteinaceous (0.528), whilst PC3 accounts for 17.6% of the variance and is charac­
matter with increasing incubation time. The remaining information terized by positive loadings of TN (0.819), NO−3 -N (0.714), and PO3− 4 -P
regarding ‘characterization of fluorescent DOM (FDOM)’ is provided in (0.722). Remarkably, temperature has become the key driver, with hot
the SI (subsection 3.1.4). spring samples exhibiting strong positive effects in response to all other
samples (Fig. 3B). This, in turn, leads to substantial changes in the
13
3.2. Changes in the biogeochemical properties of glaciers-fed specific concentrations of DSi, NH+ 4 -N, TN, DON, δ C-DIC, DOC, pH, NO3 -N,
−

ecosystems and PO3−4 -P.

Overall, FDOM components are quite similar in water samples
Biogeochemical properties of glaciers-fed specific ecosystems (Pan­ collected from different ecosystems (glaciers, streams, tributaries, the IR
gong Tso saline lake, pond, groundwater, and hot springs) are sub­ mainstream, saline lake, pond, hot springs, and groundwater), with the
stantially altered and feature typical differences among them. In exception of sewage-affected groundwater (Fig. 6, Fig. S5), thus they
particular, DOC concentration is largely different and follows the order: represent a unique signature of the glacial-fed ecosystems of UIRB. Such
saline lake (6.6–6.9 mg/L) > pond (3.1–3.2 mg/L) > hot springs uniqueness is preserved and maintained by several environmental and
(1.8–2.9 mg/L) > groundwater (1.5–1.8 mg/L), in comparison with biogeochemical functions and factors, including the sub-zero ambient
glaciers, streams, tributaries, and the IR mainstream (1.1–2.6 mg/L). AT, the extremely low winter temperature (approximately − 28 ◦ C in
Similarly, the δ13C-DIC values also vary significantly and follow the January–February), the relatively scarce precipitations, the occurrence
order: saline lake (3.3‰–3.4‰) > pond (− 9.1‰ to − 9.0‰) > ground­ of barren mountains with extremely sparse vegetation, and the complex
water (− 13.7‰ to − 11.8‰) > hot springs (− 4.0‰ to − 1.4‰), in hard-rock aquifer system that determines a relatively low microbial
comparison with glaciers, streams, tributaries, and the IR mainstream activity in the dry-arid UIRB. The biogeochemical properties of Pangong
(− 9.5‰ to − 1.5‰). Remarkably, the differences in DOC and δ13C-DIC Tso saline lake, pond, groundwater, and hot springs are further
values between glaciers and the samples of saline lake, hot springs, described in detail in the SI, subsection 3.2.
pond, and groundwater exhibit either a significant or even a highly
significant level by a two-sample z-test (Table S3). Similar wide varia­
tions are also observed in TN, DON, and nutrients. Such differences 3.3. Dynamic changes and cycling of glacier photosynthetically-derived
among specific ecosystems might be ascribed to further rapid reproc­ OM
essing/recycling of glacier-fed meltwaters by diverse processes that are
active in each system. For example, high DOC and high δ13C-DIC values The conceptual model (Fig. 7) depicts the production processes of
in the studied saline lake are the likely consequence of two simultaneous photosynthetically-derived OM in glaciers (Remias et al., 2005; Karsten
processes, i.e., substantial photosynthetic activity coupled with high and Holzinger, 2014; Smith et al., 2017), as occurring by uptake of at­
evapotranspiration under high salinity (14100–14500 μS/cm) at rela­ mospheric CO2 (δ13C-DIC, 8.4‰ in 2015, Graven et al., 2020) and fol­
tively high altitude (Cory et al., 2014; Smith et al., 2017; Bade et al., lowed by release of FDOM, DIC, and nutrients by photoinduced and/or
2004; Yi et al., 2021; Morana et al., 2015). High DOC content in pond biological respiration. The uptake of atmospheric CO2 is supported by
water is due to high primary productivity through photosynthesis the depleted δ13C-DIC (− 9.5‰) measured at the top of the Khardungla
(McKnight et al., 1994; Brown et al., 2004; Cory and McKnight, 2005) glacier, which is presumably caused by the degradation of DOM or
under conditions of relatively high WT (25ᵒC), whereas the highly
depleted δ13C-DIC value comes from newly-released lighter CO2/DIC. Table 2
The latter may derive from the degradation of DOM/FDOM originated Loadings of parameters on significant principal components and total variance
from photosynthetically-derived OM (Stedmon et al., 2007b; Stubbins explained for glaciers, ponds, hot springs, and saline lakes of UIRB, Western
Himalayas. Bold values indicate strong and moderate associations.
et al., 2010; Singer et al., 2012; Bade et al., 2004). Conversely, the
lowest DOC content in groundwater may be due to infiltration of Variables PC1 PC2 PC3 PC4 PC5
glaciers/snow-fed meltwaters, rainwater, and human-affected tubewell T C
◦
0.011 0.917 − 0.192 0.148 − 0.264
waters. The depleted δ13C-DIC values in groundwater may be ascribed to pH 0.799 − 0.392 0.067 − 0.184 0.200
fresh inputs from carbonate and silicate rock-soil bed aquifers, which EC 0.922 − 0.073 0.248 0.227 − 0.023
DOC 0.919 − 0.099 0.183 0.205 − 0.087
would occur during infiltration of the glaciers-fed meltwater and rain­ δ13C-DIC 0.892 0.119 − 0.173 0.044 − 0.198
water infiltration (Lone et al., 2021; Bhat et al., 2023). Importantly, the TN 0.026 0.495 0.819 − 0.227 0.157
relatively low-temperature hot springs (54 ◦ C) show the highest values DON 0.921 0.038 0.272 0.178 − 0.021
of TN, DON, NH+, 4 and DSi, which gradually decrease at medium tem­
NO3-N − 0.579 − 0.145 0.714 0.239 0.001
NH+4 -N 0.329 0.683 0.337 − 0.513 0.178
perature (65 ◦ C) and substantially decrease at the highest temperature
NO2-N − 0.028 0.528 − 0.220 0.655 0.481
(71ᵒC). At the same time, the δ13C-DIC values gradually enriched PO4-P − 0.377 − 0.090 0.722 0.329 − 0.235
(Table S4). These results suggest that temperature-dependent DOM DSi − 0.020 0.976 − 0.105 − 0.002 − 0.142
production and its subsequent mineralization are promoted at increas­ Eigenvalue 4.563 2.997 2.116 1.093 0.521
ingly high WT in hot springs. Variability (%) 38.026 24.972 17.629 9.104 4.342
Cumulative % 38.026 62.998 80.627 89.731 94.073
Substantial dynamic changes and/or cycling of DOC, δ13C-DIC, and
M.A. Bhat et al.

Fig. 7. Conceptual model of production processes of photosynthetically-derived primary producers (e.g. phytoplankton, algae) via photosynthesis in glaciers (Drang
Drung glacier, Ladakh) by uptake of atmospheric δ13C-DIC (− 8.4‰; Graven et al., 2020) in the presence of sunlight, and subsequent release of FDOM, CO2/DIC
(δ13CDIC:‒9.5‰ to − 0.6‰) and nutrients under photoinduced and/or biological respiration. The δ13C-DIC values for phytoplankton, glaciers, streams, tributaries,
Indus River (IR) mainstream-the upper part, IR mainstream-the lower part, saline lake, pond, groundwater and hot springs are presented in the diagram.

FDOM following its input from photosynthetically-derived PP, i.e., OM. N and imply the release of N into the atmosphere. Remarkably, C and N
In particular, the CO2 and DIC produced by DOM deriving from the transformations in the upper part of the IR mainstream might be favored
partial degradation and respiration of photosynthetic OM show the same by the relatively high WT (23.5 ± 5.1 ◦ C; Table S1) which is further
δ13C-DIC value (− 9.5‰), which is substantially enriched (− 0.6‰) by supported by the significant positive correlations of WT with DOC, NH+ 4-
the higher productivity over time (Lammers et al., 2017; Yi et al., 2021). N, and NO−2 -N (Table S4). These overall changes of C and N together
The DOC concentrations appear to increase during water transport with δ13C-DIC under the influences of WT and pH are substantially
from glaciers to streams (7.9%) and then tributaries (2.5%), followed by supported by PCA(Table 1; Fig. 3), in which PC1 and PC2 highlight the
mineralization (5.3% decrease) from streams to tributaries, along with simultaneous alterations of DOM dynamic and cycling, and N-nutrients
enrichment of δ13CDIC from –(5.3 ± 2.5‰) to –(4.4 ± 2.1‰) and then under the influence of the ambient temperature. Furtehrmore, PC3 and
–(4.3 ± 1.6‰). Such DOC changes substantially occur in the upper and PC4 emphasize the transformation of δ13C-DIC and NO−3 -N with corre­
lower parts of the IR mainstream, with an average increase of, respec­ sponding loss or degradation of DOM, NH+ 4 -N, and NO2 -N under the
−

tively, 23.4% and 17.9% compared to glaciers, along with a minerali­ influence of pH. Finally, PC5 explains 9.8% of the variance, with high
zation degree amounting to 4.5% in the lower part with respect to the positive loadings for EC (0.736) and DSi (0.486), as well as negative
upper part (Table S2). Simultaneously, δ13CDIC values are depleted in the loadings of DOC (− 0.400) and NO−2 -N (− 0.309), which highlights the
upper part of the IR mainstream (− 4.4 ± 1.0‰) compared to tributaries, relation between ionic strength of the water and DSi that might be
whilst enriched values are observed in the lower part of the IR main­ influenced by the loss/degradation of DOM and NO−2 -N. Remarkably,
stream (− 4.0 ± 1.5‰). These findings imply that organic carbon is temperature has emerged as the key driver, exhibiting a positive
constantly produced and degraded, with the production of CO2 that is response on all samples from the mainstream IR in the upper part, as
constantly emitted into the atmosphere, thereby contributing to the well as a few samples from streams, tributaries, and the mainstream IR
global carbon cycle. in the lower part (Fig. 3A). In contrast, temperature has a lesser effect
Such changes in DOC take place alongside increases in TN (by 13% across all samples from the glaciers, along with a few samples from
and 6.7%, respectively), DON (by 8.0% and 8.9%, respectively), NO−3 -N streams, tributaries, and the mainstream IR in the lower part (Fig. 3A).
(by 17% and 6.3%, respectively), and NO−2 -N (by 71% and 218%, This, in turn, influences on the production and/or mineralization of
respectively), as well as a decrease of NH+ 4 -N (by 4.5% and 8.8%, DOM, along with subsequent alterations in N- and P-nutrients. This
respectively) in streams and tributaries, compared to glaciers (Table S2). agrees with earlier studies, showing that warmer temperatures enhance
Moreover, TN, DON, and NH+ 4 -N reach their highest levels (14.5%, photosynthesis and respiration, which, in turn, indirectly promote N2-
106.8% and 13.0%, respectively), and NO−3 -N reaches its lowest level fixation by planktonic communities (Li et al., 2019; Großkopf and
(13.5%) along with an increase of NO−2 -N (104.3%) in the upper part of LaRoche, 2012; Yang et al., 2021; Inomura et al., 2019). The continuous
the IR mainstream compared to glaciers. At the same time, all of these N- enrichment of δ13C-DIC values from glaciers to streams, tributaries, and
nutrients (except NO−3 -N) decrease when passing from the upper to the the IR mainstream (lower part) would indicate the occurrence of CO2
lower part of the IR mainstream whereas NO−3 -N increases by approxi­ emissions into the atmosphere from DOM degradation, which would
mately 22.0%. These results suggest that continuous production- occur together with denitrification that additionally emits N2 to the
degradation of DON would reflect the occurrence of nitrification and atmosphere.
denitrification processes, which would account for the increase of NO−2 - Based on the average annual water discharge in the IR mainstream
M.A. Bhat et al.

(1354 m3/s) (Bhat et al., 2023; Mukhopadhyay and Dutta, 2010), during water transport, thereby contributing to the global C cycling,
flowing from glaciers via streams and tributaries and on the absence of and/or supporting downstream microbial metabolism, with an influence
other point sources (e.g. precipitation, human activities/agricultural on primary productivity in downstream locations (Hood et al., 2009;
runoff), the net C emissions as CO2 can be calculated based on DOM Singer et al., 2012). Secondly, N-nutrients are emitted as N2/N2O into
mineralization (i.e., decreasing DOC) during water transport in tribu­ the atmosphere via nitrification-denitrification processes (Van Kessel
taries and the IR mainstream (lower part) compared to, respectively, et al., 2015; Ward et al., 2009), and phosphates are converted into
tributaries and the IR mainstream (upper part). These emissions amount phosphate minerals by sedimentary deposition (Reinhard et al., 2017;
to approximately 226.5 g/s or 7.1 × 109 g/year (see detailed calcula­ Hao et al., 2022). Thirdly, both N- and P-nutrients may partly contribute
tions in Text SI 2.6). Similarly, the net N emissions based on decreasing to photosynthesis in the downward margins/downstream waters, by
TN from streams to tributaries and the IR mainstream (upper part) to the potentially fueling the microbial food web and microbial metabolism
IR mainstream (lower part) amount to approximately 61 g/s or 1.9 × (Lund et al., 2018; Tyrrell, 1999; Elser et al., 2007).
109 g/year. In contrast, the net C sequestration based on increasing DOC Increasing ambient temperature due to climate change may contin­
in streams compared to glaciers and in the IR mainstream (upper part) uously reduce the net amount and volume of ice storage in glaciers (Nie
compared to tributaries is approximately 596.88 g/s or 1.9 × 1010 et al., 2021; Rounce et al., 2023), accelerating summer meltwater
g/year. Likewise, the net N sequestration based on increasing TN in run-off, with the consequent increase of physical erosion and chemical
streams compared to glaciers and in the IR mainstream (upper part) weathering beneath glaciers (Li et al., 2022). Climatic-induced changes
compared to the IR mainstream (lower part) amounts to approximately during 2000–2020 have caused the highest glacier melt rates in the
107 g/s or 3.4 × 109 g/year. Western Himalayas (Brun et al., 2017), which, in turn, have determined
These transformation processes, i.e., the production-degradation- the shrinkage of the overall glacier cover by 5.31 ± 0.33 km2, along with
emission of DOM/FDOM, CO2/DIC, and N-nutrients from glaciers to snow retreat, thickness changes, mass loss, and velocity changes of 77
streams, tributaries, and the IR mainstream, will primarily affect the glaciers in the Drass basin, Western Himalaya (Romshoo et al., 2022).
DOM/FDOM components by two distinct pathways. Firstly, the CO2/DIC Based on the global temperature change scenarios, glaciers are projected
are the ending mineralization byproducts of DOM/FDOM, and they may to lose 26 ± 6% (+1.5 ◦ C) to 41 ± 11% (+4 ◦ C) of their mass by 2100,
involve either lighter CO2 emission into the atmosphere during water relative to 2015 (Rounce et al., 2023). Therefore, possible C and N
transport (thereby enriching the remaining δ13CO2/DIC) (Morana et al., emissions are projected to increase, respectively, to approximately 9.0
2015; Yi et al., 2021; Bade et al., 2004), or a new input of lighter × 109 g/year and 2.4 × 109 (+1.5 ◦ C) to 1.0 × 1010 g/year and 2.7 × 109
CO2/DIC from DOM/FDOM degradation, which would replace the (+4 ◦ C) over the same timescale.
depleted δ13CO2/DIC in solution (Stedmon et al., 2007b; Bhatia et al., A warmer climate is expected in UIRB in the near future, with a
2010; Stubbins et al., 2010; Singer et al., 2012; Bade et al., 2004). These temperature increase from mid-century (2041–2071) to late 21st cen­
processes are primarily responsible for the widely variable δ13C-DIC tury (2071–2100) evaluated from 2.36 ◦ C to 3.5 ◦ C under the RCP4.5
values in glaciers (− 5.3 ± 2.5‰), streams (− 4.4 ± 2.1‰), tributaries scenario, and 2.92 ◦ C–5.23 ◦ C under RCP8.5 (Shah et al., 2020). Such
(− 4.3 ± 1.6‰), and the upper and lower parts of the IR mainstream climatic changes in glaciers are expected to further alter the streamflow
(− 4.4 ± 1.0‰ and − 4.2 ± 1.5‰, respectively). Secondly, the N- and patterns and the seasonality of glaciers runoff, thereby shifting the
P-nutrients are the byproducts of photochemically and biologically snowmelt runoff as a function of the mean temperature rise in spring. In
active DON and DOP, i.e., two important biogenic components of turn, these processes will affect the incidence of glacial-lake outburst
DOM/FDOM. They are subject to continuous transformation processes floods and increase the risk of flooding in some months, while the
consisting of either new production from photosynthetically-derived decline of meltwater flow in other months might cause a drought (Kaser
OM (Stedmon et al., 2007b; Zhang et al., 2021; Yang et al., 2021), et al., 2010; Nie et al., 2021; Tahir et al., 2011; Pritchard, 2019).
which causes an increase in their concentrations during water transport, Furthermore, rainfall-induced glaciers-fed meltwaters from the Kar­
or emission of N-nutrients as N2/N2O into the atmosphere via akorum, Himalayas, and Hindu-Kush highlands, which cover 2200 km2
nitrification-denitrification processes (Van Kessel et al., 2015; Ward of permanently glaciated area (Shrestha et al., 2015), and feed the UIRB
et al., 2009). Moreover, phosphates would decrease by sedimentary system, are expected to increase substantially during winter and sum­
deposition of phosphate minerals (Reinhard et al., 2017; Hao et al., mer. Ultimately, these changes will have a negative impact on sustaining
2022). Although N- and P-nutrients would provide little contribution to downstream hydrology/ecology and aquatic organisms, thereby poten­
photosynthesis in streams, tributaries, and the IR mainstream, because tially compromising microbial metabolism by enhancing primary pro­
of the continuous water movement, they would definitely enrich the ductivity via photosynthesis, elemental cycling, out-gassing, and
microbial food web and microbial metabolism in downward margins precipitation of carbonates. Furthermore, all these processes are
and stagnant water bodies (Lund et al., 2018; Deregibus et al., 2016; modulated by various physicochemical conditions in the glacier’s
Hopwood et al., 2020). Finally, overall OM transformations would be downward margins (Jan et al., 2010; Hood et al., 2009; Singer et al.,
regulated by several environmental factors, such as the ambient WT that 2012).
varies from 8 ◦ C to 32 ◦ C in glaciers, streams, tributaries, and the IR
mainstream, the ambient AT below zero, the high temperature in the 4. Conclusion
upper part of the IR mainstream, sunlight irradiance, and the barren
mountains with extremely sparse vegetation in the dry-arid UIRB. The origin of photosynthetic OM in glaciers top-bed, the subsequent
in-situ input of DOM, and its transformation in glaciers and then in the
3.4. Environmental implications of UIRB glaciers and their downward streams-tributaries-IR mainstream system can be studied by measuring
fate in C and N cycling the corresponding changes of δ13C-DIC, DOC, FDOM, and nutrients. The
lowest δ13C-DIC value (− 9.5‰) in the glacier’s top is the key signature
The autochthonous DOM/FDOM of planktonic origin in glaciers, of the site where photosynthetic OM would originate from atmospheric
particularly top-mountain glaciers, is rapidly transformed in the CO2, which features a very similar δ13CO2 value (− 8.4‰ in 2015;
downward glacial melts and then in streams, tributaries, and the IR Graven et al., 2020). The subsequent increase of δ13C-DIC in glacier
mainstream, which suggests a high lability and capability of generating melts and the subsequent changes occurring in the downward margins
significant amounts of CO2, DIC, NO−3 -N, NH+ 4 -N, NO2 -N, PO4-P, and
−
would involve a continuous transformation of DOM/FDOM and its DON
various fluorescent compounds. These degradation byproducts of DOM components. These rapid transformations, reported to occur elsewhere,
and their downward fate are associated with several biogeochemical are primarily responsible for autochthonous DOM production from OM
phenomena. Firstly, lighter CO2/DIC is emitted into the atmosphere by rapid microbial metabolisms. Thus, they have the potential of
M.A. Bhat et al.

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