Why did cultivated apples never appear on Jifa's table?

Produced by: Science Popularization China

Author: Li Tong (Institute of Botany, Chinese Academy of Sciences)

Producer: China Science Expo

There is a behind-the-scenes clip from the recently popular movie "Fengshen". The actor who played the young emperor Jifa successfully got the role of Jifa after filming the horse riding scene because of his good performance. When saying goodbye to the white horse (he played "Snow Dragon Colt" in the movie) that he worked with, he finally fed it an apple to thank "Snow Dragon Colt" for helping him start his journey as an actor.

Ji Fa rode the Snow Dragon Horse and galloped home

(Photo source: Stills from "Investiture of the Gods")

This makes people wonder, if you travel back to the world of "Fengshen", what gift would you give to Ji Fa to make him impressed?

The answer may be apples . It's not because Jifa loves apples, but because he has never seen apples cultivated .

Although Ji Fa is a character in the myth of "Investiture of the Gods", there was a real person in history, that is King Wu of Zhou . King Wu of Zhou, Ji Fa, died around 1043 BC, and the earliest cultivated apples were introduced to China during the Yuan Dynasty (1271-1368). According to the historical era, he really had never eaten the crispy and sweet cultivated apples.

Wild apples travel far and wide

Fruits, like livestock, were domesticated by humans before they stumbled onto our tables. The many varieties of apples we eat today are cultivated apples , which are domesticated through continuous hybridization of wild apples and are "evolved" species that have been selected by human tastes for survival of the fittest .

The origin and cultivation of apples have always been a concern of modern botanists, even writers and archaeologists. The most mainstream (but not the only) view is that the ancestor of cultivated apples is the wild species of Sevastopol apples, which traveled westwards to Europe along the ancient Silk Road and hybridized with oriental apples in the Caucasus and forest apples in Eastern Europe, forming a variety of European apple varieties. European apple varieties traveled eastwards and were introduced to the mainland, where they hybridized and domesticated with wild apple species in the mainland, producing the early Chinese cotton apple, commonly known as "Lin Qin", which was then gradually cultivated.

Coincidentally, according to research by writers and agronomists, the earliest record of apples appeared in "Shanglin Fu" written by Sima Xiangru during the Western Han Dynasty (also known as the Former Han Dynasty, 202 BC - 8 AD), which recorded "So Lu oranges ripened in summer, yellow oranges and tangerines, loquats and persimmons, Tingnai (nài) Magnolia officinalis, yǐng (yǐng) dates and bayberries, cherries and Putao, hidden husbands and li (yù) li, Dada and Lizhi, spread out in the harem and lined up in the north garden."

Among them, "Houpu" refers to pear, and "Nai" refers to cotton apple . "Shanglin Fu" is what netizens often say, "Write Shanglin Fu, get your sweetheart". The Shanglin Garden described in it is near Xi'an, Shaanxi Province today, which is Xiqi during the Shang and Zhou dynasties . Perhaps this is the destined fate between the "Snow Dragon Horse" in the movie "Investiture of the Gods" and Xiqi.

Ringo's debut was a huge success

After the Tang Dynasty (618-907), nai also gave rise to lingong (nai) and nai li (nai plum). This can be verified by the description of Tang Dynasty poet Bai Juyi in "Xi Sheng Du Hua Yi Zhongzhou Dongpo New Flower Trees, So I Sent a Poem to Donglou": "I remember Dongpo's red and rotten (màn), wild peach mountain apricot water lingong (manu)" and Zhang Xiaobiao's poem: "White birds are fascinated by mountain peonies, and red-dressed prostitutes are jealous of water lingong (manu)".

Qing Dynasty, Wu Qijun's "Illustrated Catalogue of Plant Names and Realities" Lin Qin inner page

(Photo source: The Paper)

The apple (Nai) is a fruit of the genus Apple in the Rosaceae family, which is what we now call the sand apple (Huahong). The apple was loved by Emperor Gaozong of the Tang Dynasty. Its grower Wang Fangyan presented the apples he grew to Emperor Gaozong and was awarded the title of Wenlinlang (official title), so the apple is also called "Wenlin fruit".

The famous Ringo also became popular in Japan. At that time, my country's Ringo was introduced to Japan by Japanese envoys to the Tang Dynasty. Therefore, Japan later called apples "Ringo", which is called "ringo" in Japanese, following the ancient Chinese tradition.

Ringo (Bonus)

(Image source: China Plant Image Library)

So how did nai plum and ringo evolve into apples?

The name Apple first appeared

The origin of the name apple can be traced back to Tang Monk in Journey to the West. According to research, with the opening of the Silk Road in the Han Dynasty, the Eastern Jin Dynasty monk Faxian first introduced Buddhist scriptures from abroad, and later the Tang Dynasty monk Xuanzang introduced Buddhist scriptures to the Tang Dynasty. As a result, some flowers and fruits in Buddhism were widely spread, and the earliest prototype of apple, "Pinpo", came from Buddhist scriptures.

Xuanzang

(Photo source: Veer Gallery)

Therefore, it can be seen that in the Tang Dynasty, people thought that Pinang and Lingo were similar, but they still couldn't tell them apart. It is said that Pinang fruit may be a red-fleshed apple native to Xinjiang, which laid the groundwork for the name apple.

In the Song Dynasty (960-1279), the apple was widely cultivated and appreciated, and was deeply loved by people. Emperor Huizong of Song wrote in "Golden Apple Traveling in Spring Orioles": "How outstanding is its good name, and its reputation dominates the spring. In March, the birds come to play and are different from others." In addition, Zhu Yi, a poet of the Southern Song Dynasty, wrote in "Begging for Pinellia Fruits from Shuangfeng Mountain" (Part 2): "I heard that the Tathagata has Buddha's eyes, and he sent me the fruits as soon as they are ripe. From now on, I will cultivate my eyes with compassion and bloom wherever there is a bloom."

It can be analyzed that the translation of Buddhist scriptures in the Song Dynasty believed that the Pinellia fruit was "Buddha's Eye Fruit" , so the Sterculia genus of the Malvaceae family in China was attributed to the Pinellia fruit, which completely drew a clear line between Pinellia and Ringo.

If you want to understand the difference between Ringo (Sand Apple) and Sterculia sterculiar, you might as well take a look at their business cards.

Name card of lingong (sand apple) : Sand apple, a small tree of the genus Apple in the Rosaceae family. The crown of the sand apple is round; the leaves are ovate or elliptical, with very fine serrations on the edges of the leaves; the umbel racemes grow at the top of the short branches, the buds are pink, and the color fades to white with a red blush after opening; the fruit is oblate, yellow or red; the flowering period is May, and the fruiting period is July-August.

The flesh is loose, sweet, sour and fragrant, and it is not resistant to storage; after a short storage, the flesh will become sandy, hence the name sand apple. Sand apple is native to Gansu and Xinjiang in northwest China. It is a unique tree species in China and is widely distributed in the Yellow River Basin, Northeast China, Hebei, and eastern Inner Mongolia at an altitude of 50-1,300 meters.

Ringo (Bonus)

(Image source: China Plant Image Library)

Name card of Artemisia annua : Artemisia annua, also known as phoenix eye fruit, commonly known as loquat fruit, seven sister fruit, is a tree of the genus Artemisia of the Malvaceae family. Its ancient name was "pinpo", which originated from Sanskrit, originally meaning "shadow", and translated as "acacia tree". It is produced in Guangdong, southern Guangxi and other places, and is also distributed in India, Vietnam and other places.

Apple

(Image source: China Plant Image Library)

When Apple Becomes an Apple with Identity

According to historical records, the first people to eat the cultivated apples that "recognized their identity" may have been the people on horseback.

The Yuan Dynasty (1271-1368) was the first unified dynasty established by a minority ethnic group, with five generations and eleven emperors. The Yuan Dynasty had a vast territory, a prosperous commodity economy and overseas trade, and frequent diplomatic exchanges with various countries. Some improved fruits produced in Central Asia were tributed to the Yuan Empire, and apples were introduced and widely cultivated.

This is proved by the tenth poem of "Hundred Chapters of Yuan Palace Ci" written by Zhu Youdun, a playwright in the Ming Dynasty: "Xinghe West Road presents the latest, and the apples are evenly distributed. They are brought into the inner court and ranked according to their grades, and are awarded to meritorious officials at the same time." The "apple" here refers to "apple". It is called "Pingbo" because it is also a "divine fruit" tributed from the Western Regions. This delicious fruit is easily associated with Pinellia (Pinellia), but Pinellia fruit (Pinellia) was recorded in the Tang Dynasty. In order to distinguish these "divine fruits", this fruit can only be called "Pingbo" or other transliterations.

Mausoleum of Genghis Khan

(Photo source: Veer Gallery)

The word "apple" first appeared in Jia Ming's "Dietary Notes" at the end of the Yuan Dynasty, and became an official name in the Ming Dynasty (1368-1644). In the agricultural book "Fangpu·Guopu" during the Wanli period of the Ming Dynasty, the entry "apple" appeared, popularizing the name of apple from a more authoritative perspective.

Science + literature, the little apple breaks the dimensional wall of disciplines

With the development of science, botanists have proposed from a scientific perspective that cultivated apples originated in Xinjiang and provided strong evidence.

From a macro perspective, botanists have analyzed the origin of apples through diversity (ecological perspective) and traits (morphological perspective).

The so-called diversity (from an ecological perspective) refers to the fact that there are many types of "mother" apples in the place of origin, and they can all serve as the source of diversity for cultivated apples. Traits (from a morphological perspective) refer to the fact that the offspring look like the "mother" apple and have similar personalities. For example, in order to explore the origin of cultivated apples, Li Yunong's team traveled to the Caucasus and Xinjiang, and spent more than ten years looking for the "mother" for cultivated apples.

1. Polymorphism of Saves apple; 2. Saves apple in Xinjiang; 3. Polymorphism of Saves apple in Xinjiang

(Image source: Reference [4])

From a microscopic perspective, with the development of science and technology and the maturity of gene sequencing technology, scientists can satisfy their curiosity about the origin of apples by observing the tiny size of pollen and chromosomes and DNA that are invisible to the naked eye.

Palynology, karyotype analysis and molecular markers are the main methods used by scientists. Palynology refers to the study of the origin of plants through the morphology of plant pollen . The smaller the volume of plant pollen, the higher the degree of evolution. The study found that among the apple plants, the pollen volume of Xinjiang wild apple is the largest, which confirms that Xinjiang wild apple is more primitive.

Core analysis refers to the fact that polyploid groups are more evolved and more adaptable. Based on this theory, it was found that the number of chromosomes in the Saves apple is relatively primitive, which supports that the Saves apple is the original species of cultivated apple.

Molecular markers are similar to the common "paternity test". The DNA sequences of different individuals are not exactly the same. Through the molecular marker method, the DNA sequence of individuals is genetically marked, and the relationship between different varieties of apples can be inferred.

Genetic control: Apple can also DIY

As curious scientists, they always bring surprises to people. Through genome research, they have also figured out the "gene controller" that can control the sweetness and size of apples.

By analyzing the genetic characteristics of the domestication and improvement process of wild apple ancestors, ancient varieties and modern varieties, the researchers found that the change in acidity played a key role in the domestication of apple flavor quality, and the genes related to sugar content remained almost unchanged. In other words, the acidity decreased without changing the sugar content, making the apple sweeter. At the same time, the Mα1 gene, a gene that affects acidity, was also found .

Modeling the evolution of fruit size and flavor during domestication of cultivated apple

(Image source: Reference [8])

Not only that, scientists have also successfully solved the factors that affect the size of apples. Apple size is related to the expression of abscisic acid (ABA) and salicylic acid (SA) . The smaller the fruit, the higher the content of abscisic acid (ABA) and salicylic acid (SA).

Plant hormones ABA and SA, which are involved in plant cell division and elongation, accumulate to high levels in low-weight fruits

(Image source: Reference [9])

Apple is a magical fruit that has brought endless inspiration to scientists, allowing them to follow in the footsteps of apples and unlock the genetic code of plants.

References:

【1】Duan Naibin. Genomic research on the origin, evolution and domestication mechanism of cultivated apple[D]. Shandong Agricultural University, 2017.

【2】Feng Tingting, Zhou Zhiqin. Research progress on the origin of cultivated apple[J]. Journal of Fruit Science, 2007(02):199-203.

【3】Luo Guihuan. Research on the Origin of Apple[J]. Journal of Beijing Forestry University (Social Sciences Edition), 2014, 13(02): 15-25.

【4】Li Yunong. Research on the origin and evolution of apple[J]. Acta Horticulturae Sinica, 1999(04):5-12.

【5】Shen Guangbin, Ding Yanyan. Study on the time when the apple was introduced into China[J]. Agricultural Archaeology, 2014(06):255-259.

【6】Yang Xiaohong, Li Yunong, Lin Peijun, et al. Study on pollen morphology and its origin and evolution of Xinjiang wild apple Malus sieversii (Ldb.) Roem[J]. Journal of Southwest Agricultural University, 1992(01):49-54.

【7】Duan, N., Bai, Y., Sun, H. et al. Genome re-sequencing reveals the history of apple and supports a two-stage model for fruit enlargement. Nat Commun 8, 249 (2017).

【8】Liao L, Zhang W, Zhang B, et al. Unraveling a genetic roadmap for improved taste in the domesticated apple. Mol Plant. 2021;14(9):1454-1471.

【9】Lin, Q., Chen, J., Liu, X. et al. A metabolic perspective of selection for fruit quality related to apple domestication and improvement. Genome Biol 24, 95 (2023).