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\begin{document}

\title{The third dimension of alpine plant leaf traits is related to
cold-tolerance}



\author[1]{Yuan Wang}%
\author[2]{Ji Suonan}%
\author[3]{Kun Liu}%
\author[4]{Yanni Gao}%
\author[1]{Sihao Zhu}%
\author[1]{Shihua Wu}%
\author[1]{Yanyan Du}%
\author[3]{Qian Liu}%
\author[3]{Si Chen}%
\author[1]{Ning Zhao}%
\affil[1]{Lanzhou University College of Pastoral Agriculture Science and Technology}%
\affil[2]{Qinghai Normal University}%
\affil[3]{Lanzhou University}%
\affil[4]{Chinese Research Academy of Environmental Sciences}%


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  \date{June 16, 2024}


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\begin{abstract}
1. Alpine plants possess unique traits owing to their adaptation to
alpine environments. However, it remains unknown whether leaf trait
relationships of alpine plants can be captured by the two general trait
dimensions of organ size and resource economics. 2. We hypothesized
that, beyond the trait dimensions of leaf size and resource economics,
non-structured carbohydrates (NSCs) would reflect an important dimension
of cold-tolerance in alpine plants. 3. We measured 12 leaf traits
critical to leaf construction, growth, and stress resistance in 225 site
species, ranging from 7 sites along the alpine steppes to alpine meadows
on the Tibetan Plateau. We also examined the cold-tolerance of 11
species at one of these sites by estimating the lethal temperature
causing 50\% frost damage (LT\textsubscript{50}). We investigated the
variation and potential leaf trait dimensions of alpine plants and
verified whether the third leaf trait dimension is related to
cold-tolerance by analyzing the relationships of NSCs and
LT\textsubscript{50}. 4. The majority of variations in 12 leaf traits of
alpine plants were captured by three trait axes, in which leaf carbon
(LCC) and NSCs (including starch, LSC and soluble sugars, LSS) were
clustered in a new dimension (PC3) beyond leaf size and structure, and
resource economics. Although LCC, LSC and LSS all showed negative
correlations with MAT, a significant negative correlation was only found
between LSS and LT\textsubscript{50}. It indicated that PC3 was able to
reflect the cold-tolerance of alpine plants to some extent, in which LSS
was the most critical trait. 5. Thus, the storage and transformation of
NSCs under stressful conditions could reflect a dimension of long-term
metabolic adaptation and cold-tolerance, which is an extension of the
resource-utilization strategy beyond construction cost and growth. Our
study suggested that NSCs should not be ignored in leaf economic
spectrum for alpine plants.%
\end{abstract}%



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\verb`Main manuscript.docx`  available at \url{https://authorea.com/users/644479/articles/657364-the-third-dimension-of-alpine-plant-leaf-traits-is-related-to-cold-tolerance}

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