跟著城市嚮導「老臺北胃」，用味道認識臺北

很多朋友來臺北，
都會問我同一個問題：
「臺北小吃那麼多，到底該從哪裡開始吃？」
夜市裡攤位一字排開、老店藏在巷弄轉角，
看起來都很有名，卻又怕吃錯、踩雷，
結果行程走完，反而沒真正記住臺北的味道。
我常被朋友笑說是「老臺北胃」。
不是因為特別會吃，而是因為在這座城市待久了，
知道哪些味道是陪著臺北人成長的日常。
這篇文章，就是我整理的一份清單。
如果你第一次來臺北，
我會帶你從這 10 樣最具代表性的臺北小吃開始，
不追一時爆紅、不走浮誇路線，
而是讓你吃完後能真正理解
原來，這就是臺灣的小吃文化。
跟著老臺北胃走，
用最簡單的方式，
把臺北的味道，一樣一樣記在心裡。

我怎麼選出這 10 大臺北小吃？

在臺北，
你隨便走進一條夜市或老街，
都可以輕易列出 30 種以上的小吃。
所以這份清單，
不是「臺北最好吃」的排名，
 而是我站在「第一次來臺北的旅客」角度，
做的推薦。
身為一個被朋友稱作「老臺北胃」的人，
我選這 10 樣小吃時，心裡一直放著幾個原則。

一吃就知道：這就是臺灣味

燒烤、火鍋很好吃，
但換個城市、換個國家，也吃得到。
我挑的，是那種
只要一入口，就會讓人聯想到的臺灣味。
 不需要解釋太多，舌頭就能懂。

不只是好吃，而是有「臺北日常感」

臺北的小吃迷人，
不只在味道，
而在它融入生活的方式。
我在意的是：

1. 會不會出現在早餐、宵夜、下班後
2. 有沒有陪伴這座城市很久的記憶

吃完之後，你會記得臺北

最後一個標準很簡單。
如果你回到家，
還會突然想起某個味道、某碗熱湯、某個攤位的香氣
那它就值得被放進這份清單裡。

接下來的 10 樣臺北小吃，
就是我會親自帶朋友去吃的在地美食。
不趕行程、不拚數量，
而是一口一口，
慢慢認識臺北。

第 1 家：饌堂－黑金滷肉飯（雙連店）｜一碗就懂臺灣人的日常

如果只能用一道料理，
 來解釋臺灣人的日常飲食，
 那我一定會先帶你吃滷肉飯。
在臺北，滷肉飯不是什麼特別的節慶料理，
 而是從早餐、午餐到宵夜，
 默默陪著很多人長大的味道。
而在眾多滷肉飯之中，
饌堂－黑金滷肉飯（雙連店），
 我很常帶第一次來臺北的朋友造訪的一家。

為什麼第一站，我會選饌堂？
饌堂的滷肉飯，走的是**「黑金系」路線**。
滷汁顏色深、香氣厚，
卻不死鹹、不油膩。
滷肉切得細緻，
肥肉入口即化，搭配熱騰騰的白飯，
每一口都是很完整、很臺灣的味道。
對第一次吃滷肉飯的旅客來說，
這種風味夠經典、也夠穩定，
不需要太多心理準備，就能理解為什麼臺灣人這麼愛它。

不只是好吃，而是「現在的臺北感」
饌堂並不是那種躲在深巷裡的老攤，
空間乾淨、節奏俐落，
卻沒有失去滷肉飯該有的靈魂。
這也是我會推薦給旅客的原因之一：
它保留了臺灣小吃的核心味道，
同時也讓第一次來臺北的人，
吃得安心、坐得舒服。

老臺北胃的帶路小提醒
如果是第一次來：

1. 一定要點招牌黑金滷肉飯
   
2. 可以加一顆滷蛋，風味會更完整
3. 搭配簡單的小菜，就很有臺灣家常感

這不是那種吃完會驚呼「哇！」的料理，
而是會讓你在幾口之後，
慢慢理解
原來，臺灣人的日常，就是這樣被一碗飯照顧著。

地址：103臺北市大同區雙連街55號1樓

電話：0225501379

菜單：https://bio.site/ZhuanTang

第 2 家：富宏牛肉麵｜臺北深夜也醒著的一碗熱湯

如果說滷肉飯代表的是臺灣人的日常，
 那牛肉麵，
 就是很多臺北人心中最有份量的一餐。
而在臺北提到牛肉麵，
 富宏牛肉麵，
 幾乎是夜貓族、加班族、外地旅客一定會被帶去的一站。

為什麼老臺北胃會帶你來吃富宏？
富宏最讓人印象深刻的，
不是華麗裝潢，
而是那鍋永遠冒著熱氣的紅燒湯頭。
湯色濃而不混，
帶著牛骨與醬香慢慢熬出的厚度，
喝起來溫潤、不刺激，
卻會在嘴裡留下很深的記憶點。
牛肉給得大方，
燉到軟嫩卻不鬆散，
搭配彈性十足的麵條，
每一口都很直接、很臺北。

不分時間，任何時候都適合的一碗麵
富宏牛肉麵最迷人的地方，
在於它陪伴了無數個臺北的夜晚。
不管是深夜下班、看完演唱會、
或是剛抵達臺北、還沒適應時差，
這裡總有一碗熱湯在等你。
對旅客來說，
這種不用算時間、不用擔心打烊的安心感，
本身就是一種臺北特色。

老臺北胃的帶路小提醒
第一次來富宏，我會這樣點：

1. 紅燒牛肉麵是首選
   
2. 如果想吃得更過癮，可以加點牛筋或牛肚
3. 湯先喝一口原味，再視情況調整辣度

這不是精緻料理，
卻是一碗能在任何時刻撐住你的牛肉麵。
在臺北，
很多夜晚，
就是靠這樣一碗熱湯走過來的。

地址：108臺北市萬華區洛陽街67號

電話：0223713028

菜單：https://www.facebook.com/pages/富宏牛肉麵-原建宏牛肉麵/

第 3 家：士林夜市・吉彖皮蛋涼麵｜臺北夏天最有記憶點的一口清爽

如果你在夏天來到臺北，
 一定會很快發現一件事
 這座城市，真的很熱。
也正因為這樣，
 臺北的小吃世界裡，
 才會出現像「涼麵」這樣的存在。
而在士林夜市，
 吉彖皮蛋涼麵，
 就是我很常帶旅客來吃的一家。

為什麼在夜市，我會帶你吃涼麵？
很多人對夜市的印象，
都是炸物、熱湯、重口味。
但真正的臺北夜市，
其實也很懂得照顧人的胃。
吉彖的涼麵，
冰涼的麵條拌上濃郁芝麻醬，
再加上切得細緻的皮蛋，
入口的第一瞬間，
就是一種「被降溫」的感覺。
那種清爽，
不是沒味道，
而是在濃香與清涼之間取得剛剛好的平衡。

皮蛋，是靈魂，也是臺灣味的關鍵
對很多外國旅客來說，
皮蛋是既好奇、又有點猶豫的存在。
但我常說，
如果要嘗試皮蛋，
涼麵是一個非常溫柔的起點。
芝麻醬的香氣會先接住味蕾，
皮蛋的風味則在後段慢慢出現，
不衝、不嗆，
反而多了一層深度。
很多人吃完後，
都會露出那種「原來是這樣啊」的表情。

老臺北胃的帶路小提醒
第一次點吉彖皮蛋涼麵，我會建議：

1. 一定要選皮蛋款，才吃得到特色
2. 醬料先拌勻，再吃，風味會更完整
3. 如果天氣真的很熱，這一碗會救你一整晚

這不是華麗的小吃，
卻非常臺北。
在悶熱的夜晚，
站在夜市人潮裡，
吃著一碗涼麵，
你會突然明白——

原來臺北的小吃，連氣候都一起考慮進去了。

地址：111臺北市士林區基河路114號

電話：0981014155

菜單：https://www.facebook.com/profile.php?id=100064238763064

第 4 家：胖老闆誠意肉粥｜臺北人深夜最踏實的一碗粥

如果你問我，
 臺北人在深夜、下班後，
 最容易感到被安慰的食物是什麼——
 我會毫不猶豫地說：肉粥。
而提到肉粥，
 胖老闆誠意肉粥，
 就是很多老臺北人口中的那一味。

為什麼這一碗粥，會被叫做「誠意」？
胖老闆的肉粥，看起來很簡單。
白粥、肉燥、配菜，
沒有華麗擺盤，也沒有複雜作法。
但真正坐下來吃，你會發現：
這碗粥，不敷衍任何一個細節。
粥體滑順、不稀薄，
肉燥香而不膩，
搭配各式家常小菜，
一口一口吃下去，
很自然就會放慢速度。
這種味道，
不是要你驚艷，
而是要你安心。

這不是觀光小吃，而是臺北人的生活片段
胖老闆誠意肉粥，
最迷人的地方，
就是它的客人。
你會看到：

1. 剛下班的上班族
2. 熬夜後來吃一碗熱粥的人
3. 熟門熟路、點菜不用看菜單的老客人

這些畫面，
比任何裝潢都更能說明這家店在臺北的位置。
對旅客來說，
這是一個走進臺北人日常的入口。

老臺北胃的帶路小提醒
第一次來吃，我會這樣建議：

1. 肉粥一定要點，這是主角
2. 配幾樣小菜一起吃，才有完整體驗
3. 不用急，慢慢吃，這碗粥就是要你放鬆

這不是為了拍照而存在的小吃，
而是那種
**會讓人記得「那天晚上，我在臺北吃了一碗很溫暖的粥」**的味道。

地址：10491臺北市中山區長春路89-3號

電話：0913806139

菜單：https://lin.ee/xxbYZyS

第 5 家：圓環邊蚵仔煎｜夜市裡最不能缺席的臺灣經典

如果要選一道
 最常出現在旅客記憶裡的臺灣小吃，
 蚵仔煎一定排得上前幾名。
而在臺北，
 圓環邊蚵仔煎，
 就是那種很多臺北人從小吃到大的存在。

為什麼蚵仔煎，這麼能代表臺灣？
蚵仔煎的魅力，
不在於精緻，
而在於它把幾種看似簡單的食材，
煎成了一種獨特的口感。
新鮮蚵仔的海味、
雞蛋的香氣、
地瓜粉形成的滑嫩外皮，
最後再淋上甜中帶鹹的醬汁，
一口下去，
就是夜市的完整畫面。
這種味道，
很難在其他國家找到替代品。

圓環邊，吃的是記憶感
圓環邊蚵仔煎，
沒有多餘的包裝，
也不刻意迎合潮流。
它留下來的原因很簡單
味道夠穩、節奏夠快、
讓人一吃就知道「對，就是這個」。
對旅客來說，
這是一家
不需要研究、不需要比較，就能安心點蚵仔煎的地方。

老臺北胃的帶路小提醒
第一次吃蚵仔煎，我會這樣建議：

1. 趁熱吃，口感最好
   
2. 不用急著加辣，先吃原味
3. 醬汁是靈魂，別急著把它拌掉

蚵仔煎不是細嚼慢嚥的料理，
它屬於人聲鼎沸、鍋鏟作響的夜市時刻。
站在人群裡，
吃著一盤熱騰騰的蚵仔煎，
你會很清楚地感受到
這，就是臺北的夜晚。

地址：103臺北市大同區寧夏路46號

電話：0225580198

菜單：https://oystera.com.tw/menu

第 6 家：阿淑清蒸肉圓｜第一次吃肉圓，就該從這裡開始

說到臺灣小吃，
 很多人腦中一定會出現「肉圓」兩個字。
但真正吃過之後才會發現，
 肉圓，從來不只有一種樣子。
在臺北，
 阿淑清蒸肉圓，
 就是我很常拿來介紹「清蒸派肉圓」的一家。

清蒸肉圓，和你想像的不一樣
不少旅客對肉圓的第一印象，
來自油炸版本，
外皮厚、口感重。
而阿淑的清蒸肉圓，
完全是另一個方向。
外皮晶瑩、滑嫩，
帶著自然的彈性，
不油、不膩，
一入口反而顯得清爽。
內餡扎實，
豬肉香氣清楚，
搭配特製醬汁，
味道層次簡單卻很乾淨。

為什麼我會推薦給第一次來臺北的旅客？
因為這顆肉圓，
不需要適應期。
它不刺激、不厚重，
即使是第一次嘗試臺灣小吃的人，
也能輕鬆接受。
對旅客來說，
這是一顆
「吃得懂、也記得住」的肉圓。

老臺北胃的帶路小提醒
第一次來阿淑，我會這樣吃：

1. 直接點一顆清蒸肉圓，吃原味
2. 醬汁先別全部拌開，邊吃邊調整
3. 放慢速度，感受外皮的口感變化

這不是夜市裡熱鬧喧囂的料理，
而是那種
安靜地展現臺灣小吃功夫的味道。
當你吃完這顆肉圓，
會更明白一件事
臺灣小吃的魅力，
往往藏在這些細節裡。

地址：242新北市新莊區復興路一段141號

電話：0229975505

第 7 家：胡記米粉湯｜一碗最貼近臺北早晨的味道

如果說前面幾樣小吃，
 是臺北的熱鬧與記憶，
 那麼米粉湯，
 就是這座城市最真實的日常。
而在臺北，
 胡記米粉湯，
 是很多人從小吃到大的存在。

為什麼米粉湯，這麼「臺北」？
米粉湯不是重口味料理，
它靠的不是刺激，
而是一碗清澈卻有深度的湯。
胡記的湯頭，
用豬骨慢慢熬出香氣，
喝起來清爽、不油，
卻能在喉嚨留下溫度。
米粉細軟，
吸附湯汁後入口順滑，
簡單到不能再簡單，
卻正是臺北人習以為常的早晨風景。

配菜，才是這一碗的靈魂延伸
在胡記吃米粉湯，
主角雖然是湯，
但真正讓人滿足的，
往往是那些小菜。
紅燒肉、豬內臟、燙青菜，
隨意點上幾樣，
湯一口、菜一口，
就是很多臺北人記憶中的早餐組合。
對旅客來說，
這是一種
不需要解釋，就能融入的臺北生活感。

老臺北胃的帶路小提醒
第一次來胡記，我會這樣建議：

1. 一定要點米粉湯，湯先喝
2. 再配 1～2 樣小菜，體驗會完整很多
3. 這一餐適合慢慢吃，不用趕

這不是為了觀光而存在的小吃，
而是一碗
每天準時出現在臺北人生活裡的湯。
當你坐在店裡，
聽著湯勺碰撞的聲音，
你會突然感覺到——
原來，臺北的早晨，
就是從這樣一碗米粉湯開始的。

地址：106臺北市大安區大安路一段9號1樓

電話：0227212120

第 8 家：藍家割包｜一口咬下的臺灣街頭記憶

如果要選一道
 外國旅客一看到就會好奇、吃完又會記住的小吃，
 割包，一定在名單裡。
而在臺北，
 藍家割包，
 就是我很放心帶旅客來認識這道經典的一站。

割包，為什麼被叫做「臺灣漢堡」？
割包的結構其實很簡單：
鬆軟的白饅頭、
燉得入味的滷五花肉、
酸菜、花生粉、香菜。
但真正迷人的，
是這些元素組合在一起時，
形成的層次感。
肉香、甜味、鹹味、清爽度，
在一口之間同時出現，
沒有誰搶戲，
卻彼此剛好。
這種平衡感，
正是臺灣小吃很迷人的地方。

藍家割包不是走浮誇路線，
它給人的感覺很直接
就是你期待中的割包樣子。
饅頭柔軟不乾，
五花肉肥瘦比例恰到好處，
入口即化卻不膩口，
花生粉的甜香收尾，
讓整體味道非常完整。
對第一次吃割包的旅客來說，
這是一個
不會出錯、也很容易愛上的版本。

老臺北胃的帶路小提醒
第一次吃藍家割包，我會這樣建議：

1. 直接點招牌割包，不要改配料
2. 如果有香菜，建議保留，味道會更完整
3. 趁熱吃，饅頭口感最好

割包不是精緻料理，
卻非常有記憶點。
站在街頭，
拿著一顆熱騰騰的割包，
邊走邊吃，
你會很清楚地感受到
這一口，就是臺灣的街頭生活。

地址：100臺北市中正區羅斯福路三段316巷8弄3號

電話：0223682060

菜單：https://instagram.com/lan_jia_gua_bao?utm_medium=copy_link

第 9 家：御品元冰火湯圓｜臺北夜晚最溫柔的一碗甜

吃了一整天的臺北小吃，
 到了這個時候，
 胃其實已經差不多滿了。
但只要天氣一涼，
 或夜色慢慢降下來，
 你還是會想找一碗——
 不是為了吃飽，而是為了舒服的甜點。
這時候，我通常會帶你來 御品元冰火湯圓。

為什麼叫「冰火」？這碗湯圓的關鍵就在這裡
御品元最有特色的地方，
就在於它的「冰火交錯」。
熱騰騰的湯圓，
外皮軟糯、內餡濃香，
搭配冰涼清甜的桂花蜜湯，
一口下去，
溫度在嘴裡交替出現。
不是衝突，
而是一種很細膩的平衡。
這樣的吃法，
也正是臺灣甜點很擅長的地方——
不張揚，但很有記憶點。

這是一碗，會讓人慢下來的甜點
和夜市裡熱鬧的甜品不同，
御品元的冰火湯圓，
更像是一個讓人停下腳步的存在。
你會發現，
坐在這裡吃湯圓的人，
說話聲都會不自覺地變小。
對旅客來說，
這不只是吃甜點，
而是一個
把白天的熱鬧慢慢收進回憶裡的時刻。

老臺北胃的帶路小提醒
第一次吃御品元，我會這樣建議：

1. 點招牌冰火湯圓，體驗完整特色
2. 先單吃湯圓，再搭配湯一起吃
3. 放慢速度，這一碗不適合趕時間

這不是為了拍照而存在的甜點，
而是一碗
會讓你記得「那天晚上在臺北，很舒服」的湯圓。

地址：106臺北市大安區通化街39巷50弄31號

電話：0955861816

菜單：https://instagram.com/lan_jia_gua_bao

第 10 家：頃刻間綠豆沙牛奶專賣店｜把臺北的味道，留在最後一口清甜

走到這一站，
 其實已經不需要再吃什麼大份量的東西了。
這時候，
 最適合的，
 是一杯不吵鬧、不張揚，
 卻會默默留在記憶裡的飲品。
頃刻間綠豆沙牛奶，
 就是我很常用來替一天畫下句點的選擇。

綠豆沙牛奶，為什麼這麼「臺灣」？
在臺灣，
飲料不只是解渴，
而是一種生活節奏。
綠豆沙牛奶看起來簡單，
但真正好喝的版本，
靠的是火候、比例，
還有耐心。
頃刻間的綠豆沙，
口感細緻、不粗顆，
甜度自然、不膩口，
牛奶的加入，
讓整杯變得柔順而溫和。
這不是衝擊味蕾的飲料，
而是一種
喝完之後，會覺得剛剛那一刻很舒服的甜。

為什麼我會用它當作最後一站？
因為它很臺北。
你可以外帶，
邊走邊喝；
也可以站在店門口，
慢慢把杯子喝空。
沒有儀式感，
卻很真實。
對旅客來說，
這杯綠豆沙牛奶，
就像是把今天吃過的所有味道，
溫柔地整理好，
帶走。

老臺北胃的帶路小提醒
第一次喝頃刻間，我會這樣建議：

1. 直接點招牌綠豆沙牛奶
   
2. 正常甜就很剛好，不用特別調整
3. 找個角落慢慢喝，別急著趕路

這一杯，
不會讓你驚呼，
卻會在回程的路上，
突然想起來。
原來，臺北的味道，是這樣結束一天的。

地址：111臺北市士林區小北街1號

電話：0228818619

菜單：https://instagram.com/chill_out_moment?igshid=YmMyMTA2M2Y=

如果只有 3 天的自助旅行在臺北，怎麼吃這 10 家？

第一次來臺北，
時間有限、胃容量也有限，
與其每一家都趕，不如照著節奏吃。
這份 3 天小吃路線，
是老臺北胃會帶朋友實際走的版本：
不爆走、不硬塞，
讓你每天都吃得剛剛好。

臺北 3 天小吃推薦行程表（老臺北胃版本）

天數	時段	店家名稱	小吃類型
Day 1	午餐	饌堂－黑金滷肉飯（雙連店）	滷肉飯
Day 1	下午	阿淑清蒸肉圓	肉圓
Day 1	晚餐	富宏牛肉麵	牛肉麵
Day 1	宵夜	胖老闆誠意肉粥	粥品
Day 2	早餐	胡記米粉湯	米粉湯
Day 2	下午	藍家割包	割包
Day 2	晚上	士林夜市－吉彖皮蛋涼麵	涼麵
Day 2	夜市	圓環邊蚵仔煎	蚵仔煎
Day 3	下午	御品元冰火湯圓	甜點
Day 3	收尾	頃刻間綠豆沙牛奶專賣店	飲品

雖然每個小吃的地點都有一點距離，但是你也知道，好吃的小吃，是值得你花一點時間前往品嘗
老臺北胃的小提醒

1. 不需要每一家都點到最滿
2. 留一點餘裕，才會想再回來
3. 臺北小吃的魅力，不在於吃多少，而在於記住了什麼味道
   

當你照著這 3 天走完，
你會發現，
臺北不是靠一兩道名菜被記住的，
而是靠這些看似日常、卻很真實的小吃。
下次再來，老臺北胃再帶你吃更深的那一輪。

老臺北胃帶路｜這 10 口，就是我心中的臺北

寫到這裡，
 其實已經不是在推薦哪一家小吃了。
而是在回頭看，
 這座城市，是怎麼用食物陪著人生活的。
滷肉飯、牛肉麵、肉粥、米粉湯，
 不是為了成為觀光名單而存在，
 而是每天默默出現在臺北人的日子裡。
夜市裡的蚵仔煎、涼麵、割包，
 熱鬧、吵雜、節奏很快，
 卻也正是臺北最真實的樣子。
而最後那碗湯圓、那杯綠豆沙牛奶，
 則是在一天結束時，
 替味蕾留下一個溫柔的句點。

如果你問我，
「這 10 家是不是臺北最好吃的小吃？」
我會說，
它們不一定是排行榜第一名，
卻是我真的會帶朋友去吃的版本。
因為它們吃得到：

1. 臺北人的日常
2. 巷弄裡的熟悉感
3. 不需要解釋，就能被理解的味道

如果你是第一次來臺北，
跟著這份清單走，
你不一定會吃得最飽，
但你一定會記得——
臺北，是什麼味道。
而如果有一天，
你又再回到這座城市，
走進熟悉的街口、
看到冒著熱氣的小攤，
你也會開始懂得，
為什麼老臺北胃，
總是記得這些看似平凡的滋味。
因為，真正留在心裡的，
從來不是吃過多少，
而是哪一口，讓你想起臺北。

胡記米粉湯不點會後悔嗎？

走完這 10 家，

你可能會發現一件事頃刻間綠豆沙牛奶專賣店吃起來順口嗎？

臺北的小吃，其實不急著被你記住。

它們就安靜地存在在街角、夜市、轉彎處，藍家割包在地人怎麼說？

等你有一天，再回到這座城市。富宏牛肉麵推薦必點嗎？

如果你是第一次來臺北，頃刻間綠豆沙牛奶專賣店不排隊會可惜嗎？

希望這份「老臺北胃帶路」的清單，

能幫你少一點猶豫、多一點安心。

不用擔心踩雷，士林夜市－吉彖皮蛋涼麵不點會後悔嗎？

也不用為了排行而奔波，圓環邊蚵仔煎值得專程去嗎？

只要照著節奏走，

你就會吃到屬於自己的臺北味道。

而如果你已經來過臺北，

那更希望這篇文章，饌堂－黑金滷肉飯（雙連店）不點會後悔嗎？

能帶你走進那些

你可能錯過、卻一直都在的日常小吃。

因為真正迷人的旅行，

從來不是把清單全部打勾，

而是某一天，

你突然想起那碗飯、那口湯、那杯甜，饌堂－黑金滷肉飯（雙連店）一定要點嗎？

然後在心裡對自己說一句：頃刻間綠豆沙牛奶專賣店本地人會吃嗎？

「下次再去臺北，還想再吃一次。」

把這篇文章存起來、分享給一起旅行的人，

或是在規劃行程時，再回來看看。

讓味道，成為你認識臺北的方式。

下一次來臺北，

別急著走遠。

老臺北胃，御品元冰火湯圓值得排隊嗎？

會一直在這些地方，

等你再回來。

Researchers discovered how valproic acid, a medication commonly used to treat epilepsy, migraines, and bipolar disorder, causes birth defects when taken during pregnancy. Researchers determined that valproic acid prevents nervous system cells from properly developing and dividing When used during pregnancy, the drug valproic acid, which is used to treat bipolar disorder, migraines, and epilepsy, can lead to birth defects. Now, research recently published in the journal PLoS Biology by Bill Keyes of the Institute of Genetics and Molecular and Cellular Biology, France, and associates gives one explanation for why: Valproic acid (VPA) causes certain nervous system development cells to enter a condition known as senescence, which prevents them from properly growing and dividing. VPA is frequently used to treat a variety of diseases. However, since its first use, there have been many instances of pregnant women using VPA giving birth to kids who had birth abnormalities such as spina bifida, facial changes, and heart malformations. A third of exposed newborns also develop cognitive decline and Autism Spectrum Disorder. Three mouse embryos, representative of the study that describes how the teratogenic drug Valproic acid can cause neurodevelopmental birth defects in mice, including microcephaly and exencephaly. The embryo on the left is a normal embryo, with no exposure to Valproic acid. The embryo in the middle is smaller and has microcephaly, while the embryo on the right exhibits exencephaly. The middle embryo and the one on the right were both exposed to Valproic acid. Credit: Muriel Rhinn (CC-BY 4.0) Role of p19Arf in VPA-Induced Developmental Defects Keyes and colleagues examined embryonic exposure to VPA in the new study by using both human organoids—three-dimensional collections of human cells generated in the lab—and mice. They found that neuroepithelial cells, which are the stem cells that give rise to the central nervous system, undergo cellular senescence as a result of VPA. The researchers also identified p19Arf as the specific molecule that caused this VPA-induced senescence. Although VPA exposure during pregnancy still resulted in other abnormalities, the scientists found that it no longer produced microcephaly (a small head size) or alterations to gene expression patterns linked to autism spectrum disorder in mice missing the p19Arf gene. Valproic acid is used to treat the manic phase of bipolar disorder, seizures, and migraine headaches. This prescription medication goes by various brand names including Depakene, Depakote, Depakote DR, Depakote ER, Depakote Sprinkles, Stavzor, and Alti-Valproic. The work is one of the first to associate cellular senescence with developmental defects, the authors say. “Overall, the discovery that atypical activation of senescence in the embryo can perturb development raises the intriguing possibility that it may also contribute to defects in developmental contexts beyond those we studied here.” Muriel Rhinn, the first author of the study, adds, “While cellular senescence has long been associated with aging and age-related disease, we now show that aberrant induction of senescence can also contribute to developmental defects. As valproic acid is strongly linked to cognitive defects and Autism Spectrum Disorder, this study now introduces an exciting link with senescence, supporting how additional studies are needed.” This study was funded by grants from La Fondation pour la Recherche Medicale (FRM) (AJE20160635985), Fondation ARC pour la Recherche sur le Cancer (PJA20181208104), IDEX Attractivité – University of Strasbourg (IDEX2017), La Fondation Schlumberger pour l’Education et la Recherche FSER 19 (Year 2018)/FRM, Agence Nationale de la Recherche (ANR) (ANR-19-CE13-0023-03) and Ligue Contre le Cancer (all to W.M.K.). I.Z.B. was supported by a 4th-year fellowship from the Fondation ARC pour la Recherche sur le Cancer and a Ph.D. fellowship from INSERM and Conseil Regional Grand-Est. A.K. was supported by a fellowship from Eur IMCBiO. The work was also supported by an institutional grant to the IGBMC, ANR-10-LABX-0030-INRT, a French State fund managed by the Agence Nationale de la Recherche under the frame program Investissements d’Avenir ANR-10-IDEX-0002-02. Sequencing was performed by the GenomEast platform, a member of the “France Génomique” consortium (ANR-10-INBS-0009). The funders had no role in the study’s design, data collection, and analysis, decision to publish, or preparation of the manuscript. Reference: “Aberrant induction of p19Arf-mediated cellular senescence contributes to neurodevelopmental defects” by Muriel Rhinn, Irene Zapata-Bodalo, Annabelle Klein, Jean-Luc Plassat, Tania Knauer-Meyer and William M. Keyes, 14 June 2022, PLoS Biology. DOI: 10.1371/journal.pbio.3001664

The SpatioTemporal Omics Consortium (STOC) is a collaborative research initiative that aims to accelerate our understanding of cellular complexity and interactions at tissue scale in development, physiology, and disease through large-scale spatially resolved multiomics analyses. International scientific consortium, the SpatioTemporal Omics Consortium (STOC), produces the first spatiotemporal maps of cellular dynamics in mice, Drosophila, zebrafish, and Arabidopsis, using BGI-Research’s world-leading Stereo-seq technology. An international team of scientists led by China’s BGI-Research published state-of-the-art panoramic spatial atlases of life, examining the cellular dynamics of organisms at different developmental stages and providing potentially significant new information for disease treatment, development, and aging, and an improved understanding of biological evolution. In a series of studies published in Cell Press journals, STOC members used the spatially resolved transcriptomics technology Stereo-seq, developed by BGI-Research, to produce spatiotemporal cellular maps of mice, small fruit flies (Drosophila), zebrafish, and the Arabidopsis plant. The papers demonstrate how Stereo-seq has achieved a major breakthrough in spatial resolution and panoramic field of view, enabling analysis of the distribution and placement of molecules and cells in situ, and over time. The paper, Spatiotemporal transcriptomic atlas of mouse organogenesis using DNA nanoball-patterned arrays, is published in Cell. The other three studies on Drosophila, zebrafish, and Arabidopsis are published in Developmental Cell. Identifying the characteristics of specific cells within a tissue has significant applications for understanding which cells are causes or indicators of disease, potentially leading to future gains in human disease research. “In the past, it took thousands or even tens of thousands of experiments to complete a spatiotemporal map. Now, with Stereo-seq developed by our scientists, it can be achieved quickly and comprehensively with one. This is a milestone breakthrough in life sciences technology advancement,” said Dr. Chen Ao, who led the development of the Stereo-seq technology at BGI-Research and is the first author of the mouse spatiotemporal atlas paper. More than 80 scientists from 16 countries, including scientists from Harvard University, Massachusetts Institute of Technology, Oxford University, University of Cambridge, the University of Western Australia, the Karolinska Institutet, the Genome Institute of Singapore, and BGI, have so far collaborated as part of STOC, an open scientific collaboration consortium focused on using spatially resolved, cellular resolution omics technologies to map and understand life. Graphic of early embryonic development of mice from 9.5 to 16.5 days using Stereo-seq technology. Credit: BGI Group The Power of Stereo-seq Technology in Understanding Cells Spatial transcriptomics technology is an emerging technology that resolves previous issues identifying characteristics of single cells within a biological tissue. It builds on the achievements of single-cell sequencing, elevating it to the next level by enabling scientists to track a cell’s precise location and how it interacts with its neighbors. To achieve this, BGI’s own sequencing patented DNA nanoball technology, which amplifies small fragments of DNA into larger samples, was combined with its in situ RNA capture technology to create Stereo-seq (SpaTial Enhanced REsolution Omics-sequencing), capable of achieving a subcellular resolution of 500 nanometers (equivalent to 0.0000005 meters) combined with a panoramic centimeter-level field of view. “The development of single-cell analytical approach over the past twenty years has really made a remarkable difference in our ability to understand how cells differ from each other. More recently, it started to be possible to combine that analysis with where cells are in a tissue or organoid tissue section,” said Patrick Maxwell, Regius Professor of Physic and Head of the School of Clinical Medicine at Cambridge, and co-author of the mouse spatiotemporal atlas paper. “In my view, this new paper takes this to a new level by combining a substantial size field of view, making it possible to analyze a tissue on a scale of a developing mouse embryo, together with a very high resolution with a very deep transcriptomic read depth.” “This enables us and the users of this data, which will be freely available, to really start to understand some very fascinating questions about how mammalian development works, and how tissues are organized. That will give us insights into developmental processes, normal tissue function, and also diseases,” he added. If we compare the study of cells to the study of the ecosystem, previous technologies allowed scientists to understand which animals or plants are on Earth. With Stereo-seq, scientists can understand which country, which area, which habitat, and which community all animals or plants belong to. At the same time, scientists can also understand what each animal is doing, their past, family history, interaction with other herds, and how they may proliferate and develop. Insights into Mouse Organogenesis and Single-Cell Mapping The scientists used Stereo-seq to examine the early embryonic development of mice, in particular from 9.5 to 16.5 days during which embryonic development is occurring at a fast rate. Stereo-seq generated the Mouse Organogenesis Spatiotemporal Transcriptomic Atlas (MOSTA), which maps the kinetics and directionality of transcriptional variation during mouse organogenesis with single-cell resolution and high sensitivity. “Stereo-seq is a transformational breakthrough in spatial transcriptomics technology and is the most powerful technology in this field of life sciences today,” said Dr. Liu Longqi of BGI-Research, one of the corresponding authors of the papers. “We now have a technology to map a panoramic atlas of every cell in an organism, according to their individual biomolecular profiles, in space and over time. We have demonstrated its robustness, and successfully mapped both animal and plant molecular physiology at a scale and resolution never before possible.” For the first time the scientists were able to produce a series of high-definition maps showing the precise location of the roughly 300,000 cells from the day 16.5 embryo. BGI-Research used this information to produce a panoramic atlas of the mouse and gain insight into the molecular basis of cell variation and differentiation in developing tissues of the brain, including the dorsal midbrain. Applications of Stereo-seq for Disease Research and Crop Breeding “The successful application of our Stereo-seq technology for development has significant implications for the future of genomic research on human diseases,” said co-corresponding author Dr. Xu Xun, director of BGI-Research. “Demonstrating that this technology can pinpoint certain cells which indicate future disease will be critical for diagnostics and therapeutics for a number of conditions.” For example, Robinow syndrome is a common birth defect. A gene related to this has been found clinically, but how this gene causes defects including cleft lip and palate, and limb shortness, is unknown. The researchers mapped the cleft lip and palate-related gene in the process of mouse embryonic development, and found that the gene was present in the lips and toes of the mouse, and showed high expression. This demonstrated that the gene is very important in the development of lips and toes in mice. If this gene is mutated, the development of lips and toes will be abnormal. This knowledge will potentially help researchers studying Robinow syndrome birth defects in humans. The BGI-led team carried out similar embryonic research with the zebrafish which has a gestation period of only 24 hours, and also produced a 3D model of the cellular map of the small fruit fly Drosophila. The spatiotemporal transcriptomic atlas of embryonic development in Drosophila, zebrafish, and mouse has opened new doors for the study of embryonic patterning and related molecular mechanisms during embryonic development, providing important data references for further work as well as a benchmark for unraveling embryonic evolution. By applying Stereo-seq research on the leaf cells of the Arabidopsis plant, the researchers were able to overcome the long-term difficulty for researchers to conduct spatially resolved single-cell omics studies on leaves and other plant tissues. BGI-Research was able to demonstrate that Stereo-seq technology can be applied in plant scientific research and crop breeding research. Some key applications include understanding key genes involved in seed development, mechanisms behind drought resistance, mechanisms behind heat resistance, and mechanisms behind salt tolerance, for staple crops from rice to wheat to maize. This could contribute to the cultivation of high-quality, stress-resistant crop strains important for many global sustainability initiatives. The studies obtained all necessary ethical clearance before they were conducted. References: “Spatiotemporal transcriptomic atlas of mouse organogenesis using DNA nanoball-patterned arrays” by Ao Chen, Sha Liao, Mengnan Cheng, Kailong Ma, Liang Wu, Yiwei Lai, Xiaojie Qiu, Jin Yang, Jiangshan Xu, Shijie Hao, Xin Wang, Huifang Lu, Xi Chen, Xing Liu, Xin Huang, Zhao Li, Yan Hong, Yujia Jiang, Jian Peng, Shuai Liu, Mengzhe Shen, Chuanyu Liu, Quanshui Li, Yue Yuan, Xiaoyu Wei, Huiwen Zheng, Weimin Feng, Zhifeng Wang, Yang Liu, Zhaohui Wang, Yunzhi Yang, Haitao Xiang, Lei Han, Baoming Qin, Pengcheng Guo, Guangyao Lai, Pura Muñoz-Cánoves, Patrick H. Maxwell, Jean Paul Thiery, Qing-Feng Wu, Fuxiang Zhao, Bichao Chen, Mei Li, Xi Dai, Shuai Wang, Haoyan Kuang, Junhou Hui, Liqun Wang, Ji-Feng Fei, Ou Wang, Xiaofeng Wei, Haorong Lu, Bo Wang, Shiping Liu, Ying Gu, Ming Ni, Wenwei Zhang, Feng Mu, Ye Yin, Huanming Yang, Michael Lisby, Richard J. Cornall, Jan Mulder, Mathias Uhlén, Miguel A. Esteban, Yuxiang Li, Longqi Liu, Xun Xu and Jian Wang, 4 May 2022, Cell. DOI: 10.1016/j.cell.2022.04.003 “Spatiotemporal mapping of gene expression landscapes and developmental trajectories during zebrafish embryogenesis” by Chang Liu, Rui Li, Young Li, Xiumei Lin, Kaichen Zhao, Qun Liu, Shuowen Wang, Xueqian Yang, Xuyang Shi, Yuting Ma, Chenyu Pei, Hui Wang, Wendai Bao, Junhou Hui, Tao Yang, Zhicheng Xu, Tingting Lai, Michael Arman Berberoglu, Sunil Kumar Sahu, Miguel A. Esteban, Kailong Ma, Guangyi Fan, Yuxiang Li, Shiping Liu, Ao Chen, Xun Xu, Zhiqiang Dong and Longqi Liu, 4 May 2022, Developmental Cell. DOI: 10.1016/j.devcel.2022.04.009 “The single-cell stereo-seq reveals region-specific cell subtypes and transcriptome profiling in Arabidopsis leaves” by Keke Xia, Hai-Xi Sun, Jie Li, Jiming Li, Yu Zhao, Lichuan Chen, Chao Qin, Ruiying Chen, Zhiyong Chen, Guangyu Liu, Ruilian Yin, Bangbang Mu, Xiaojuan Wang, Mengyuan Xu, Xinyue Li, Peisi Yuan, Yixin Qiao, Shijie Hao, Jing Wang, Qing Xie, Jiangshan Xu, Shiping Liu, Yuxiang Li, Ao Chen, Longqi Liu, Ye Yin, Huanming Yang, Jian Wang, Ying Gu and Xun Xu, 4 May 2022, Developmental Cell. DOI: 10.1016/j.devcel.2022.04.011 “High-resolution 3D spatiotemporal transcriptomic maps of developing Drosophila embryos and larvae” by Mingyue Wang, Qinan Hu, Tianhang Lv, Yuhang Wang, Qing Lan, Rong Xiang, Zhencheng Tu, Yanrong Wei, Kai Han, Chang Shi, Junfu Guo, Chao Liu, Tao Yang, Wensi Du, Yanru An, Mengnan Cheng, Jiangshan Xu, Haorong Lu, Wangsheng Li, Shaofang Zhang, Ao Chen, Wei Chen, Yuxiang Li, Xiaoshan Wang, Xun Xu, Yuhui Hu and Longqi Liu, 4 May 2022, Developmental Cell. DOI: 10.1016/j.devcel.2022.04.006

This rotifer has just survived a life-threatening infection. When a fungal disease attacked, she switched on hundreds of genes that her ancestors copied from microbes, including antibiotic recipes stolen from bacteria. Credit: C. G. Wilson 2019 Bdelloid rotifers, a type of small freshwater animal, harness stolen bacterial genes to create antibiotics, offering insights into developing safer antimicrobial drugs and addressing growing antibiotic resistance. A team of researchers from the University of Oxford, the University of Stirling, and the Marine Biological Laboratory (MBL), Woods Hole discovered that a group of tiny, freshwater animals protect themselves from infections using antibiotic recipes “stolen” from bacteria. These microscopic creatures are called bdelloid rotifers, which means ‘crawling wheel-animals’. Although they are smaller than a hair’s breadth, they have a head, mouth, gut, muscles, and nerves like other animals. Genetic Defense Mechanisms The study, recently published in Nature Communications, reveals that when these rotifers are exposed to fungal infection, they activate hundreds of genes that they acquired from bacteria and other microbes. Some of these genes produce resistance weapons, such as antibiotics and other antimicrobial agents, in the rotifers. “When we translated the DNA code to see what the stolen genes were doing, we had a surprise,” said lead study author Chris Wilson of the University of Oxford. “The main genes were instructions for chemicals that we didn’t think animals could make — they looked like recipes for antibiotics.” Like other animals, bdelloid rotifers need strategies to fight off infections and avoid ending up like this diseased individual, which has been taken over and killed by a fungus. Credit: C. G. Wilson 2024 Prior research found that rotifers have been picking up DNA from their surroundings for millions of years, but the new study is the first to discover them using these genes against diseases. No other animals are known to “steal” genes from microbes on such a large scale. “These complex genes – some of which aren’t found in any other animals – were acquired from bacteria but have undergone evolution in rotifers,” said study co-author David Mark Welch, senior scientist and director of the Josephine Bay Paul Center at the Marine Biological Laboratory. “This raises the potential that rotifers are producing novel antimicrobials that may be less toxic to animals, including humans, than those we develop from bacteria and fungi.” Unveiling Unique Antibiotic Production Antibiotics are essential to modern healthcare, but most of them were not invented by scientists. Instead, they are produced naturally by fungi and bacteria in the wild, and humans can make artificial versions to use as medicine. The new study suggests that rotifers might be doing something similar. “These strange little animals have copied the DNA that tells microbes how to make antibiotics,” explains Wilson. “We watched them using one of these genes against a disease caused by a fungus, and the animals that survived the infection were producing 10 times more of the chemical recipe than the ones that died, indicating that it helps to suppress the disease.” This bdelloid rotifer is using her wheels to hoover up and eat bacteria and other particles floating in the water. This animal is about half a millimeter long, and the wheels are on the head end. Credit: C. G. Wilson 2024 Implications for New Antibiotics The scientists think that rotifers could give important clues in the hunt for drugs to treat human infections caused by bacteria or fungi. Antibiotics are becoming less effective because the disease-causing microbes have evolved to become resistant and no longer respond to treatment. The World Health Organization recently sounded the alarm, warning in a June report of the “pressing need” to develop new antibiotics to counter the threat of resistance. “The recipes the rotifers are using look different from known genes in microbes,” said study author Reuben Nowell of the University of Stirling. “They’re just as long and complicated, but parts of the DNA code have changed. We think the recipe has been altered by a process of evolution to make new and different chemicals in the rotifers. That’s exciting because it might suggest ideas for future medicines.” The genes the rotifers acquired from bacteria encode an unusual class of enzymes that assemble amino acids into small molecules called non-ribosomal peptides. “The next phase of this research should involve identification of multiple non-ribosomally synthesized peptides produced by bdelloid rotifers, and establishment of the conditions upon which the synthesis of these compounds can be induced,” said study co-author Irina Arkhipova, senior scientist at the Marine Biological Laboratory. One problem with developing new drugs is that many antibiotic chemicals made by bacteria and fungi are poisonous or have side effects in animals. Only a few can be turned into treatments that clear harmful microbes from the human body. If rotifers are already making similar chemicals in their own cells, they could lead the way to drugs that are safer to use in other animals, including people. Understanding Rotifer Gene Acquisition A big question is why rotifers are the only animals that borrow these useful genes from microbes at such high rates. “We think it might be linked with another strange fact about these rotifers,” said Tim Barraclough, a study co-author from the University of Oxford. “Unlike other animals, we never see male rotifers. Rotifer mothers lay eggs that hatch into genetic copies of themselves, without needing sex or fertilization.” According to one theory, animals that copy themselves like this can become so similar that it starts to be unhealthy. “If one catches a disease, so will the rest,” explained Barraclough. Because bdelloid rotifers don’t have sex, which allows the parental genes to recombine in beneficial ways, the rotifer mother’s genome is directly transferred to her offspring without introducing any new variation. “If rotifers don’t find a way to change their genes, they could go extinct. This might help explain why these rotifers have borrowed so many genes from other places, especially anything that helps them cope with infections,” said Barraclough. Nowell thinks there is much more to learn from rotifers and their stolen DNA “The rotifers were using hundreds of genes that aren’t seen in other animals. The antibiotic recipes are exciting, and some other genes even look like they’ve been taken from plants. The findings are part of a growing story about how and why genes get moved between different kinds of life,” he said. Reference: “Bdelloid rotifers deploy horizontally acquired biosynthetic genes against a fungal pathogen” by Reuben W. Nowell, Fernando Rodriguez, Bette J. Hecox-Lea, David B. Mark Welch, Irina R. Arkhipova, Timothy G. Barraclough and Christopher G. Wilson, 18 July 2024, Nature Communications. DOI: 10.1038/s41467-024-49919-1

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