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。

饌堂－黑金滷肉飯（雙連店）值得排隊嗎？

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需要加料嗎？

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，頃刻間綠豆沙牛奶專賣店不排隊會可惜嗎？

等你有一天，再回到這座城市。阿淑清蒸肉圓推薦點什麼？

如果你是第一次來臺北，胖老闆誠意肉粥吃過會想再來嗎？

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，饌堂－黑金滷肉飯（雙連店）推薦點什麼？

也不用為了排行而奔波，御品元冰火湯圓女生會喜歡嗎？

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，阿淑清蒸肉圓會不會太甜？

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，圓環邊蚵仔煎當正餐適合嗎？

然後在心裡對自己說一句：圓環邊蚵仔煎真的好吃嗎？

「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，圓環邊蚵仔煎會踩雷嗎？

會一直在這些地方，

等你再回來。

A mouse brain section highlighting the hippocampus is overlaid with the molecular structures of the anesthetics isoflurane (purple), medetomidine/midazolam/fentanyl (orange), and ketamine/xylazine (red). The four panels in the lower part show two-dimensional maps of active neurons in the same hippocampal region during wakefulness (top left) and under the different anesthetics indicated above. Colors indicate the respective anesthetics. Neurons with correlated activity are connected by black lines. Credit: Simon Wiegert, CC-BY Among typical anesthetics, only isoflurane results in undisrupted memory formation in mice. Memory loss is common after general anesthesia, particularly for events occurring immediately before surgery — a phenomenon called retrograde amnesia. But a new study published on April 1st, 2021, in the open access journal PLOS Biology, led by Simon Wiegert at the University Medical Center Hamburg-Eppendorf in Germany, shows that changes in the hippocampus — the part of the brain used to make new memories — differ depending on which general anesthetic is used. Consequently, their effects on memory formation also differ. Understanding how different anesthetics affect the brain, particularly the hippocampus, is therefore important for both clinicians with human patients and experimental scientists who work with animals. Wiegert and his team recorded brain activity from the hippocampus while mice were anesthetized using one of three common combinations of general anesthetics: isoflurane, ketamine/xylazine (Keta/Xyl), and medetomidine/midazolam/fentanyl (MMF). Brain activity was recorded electrically and by optically imaging calcium flow — a dynamic signal indirectly reflecting neuronal activity and a trigger that allows brain cells to pass signals to each other. Both recording methods showed that each drug changed brain activity in the hippocampus compared to wakefulness or natural sleep. The researchers found a number of differences in how the specific anesthetics affected the brain. For example, Keta/Xyl strongly reduced overall calcium activity, while MMF affected its rate much more than the duration. Furthermore, all anesthetics affected the stability of synaptic connections between brain cells in the hippocampus. Keta/Xyl disturbed synaptic stability most drastically, reflecting its strong disturbances of neuronal calcium activity. Recovery time also differed; brain activity returned to normal in about 45 minutes after isoflurane anesthesia, but it took close to 6 hours for the other two drugs. Similarly, the mice showed signs of retrograde amnesia after both Keta/Xyl and MMF anesthesia. But after isoflurane anesthesia — the condition, which showed the mildest disturbances compared to natural sleep — they could still remember what they had learned before the surgery. Knowing these varying effects on the hippocampus and memory formation should be useful for doctors or experimenters when considering which method to use. Dr. Wiegert notes, “200 million general anesthesia are administered worldwide per year, and amnesia is a central part of it. So, we wondered how it affects the (mouse) hippocampus — a brain region essential for the formation of everyday memories. Surprisingly, the literature is scarce and inconsistent. Therefore, we teamed up with the group of Ileana Hanganu-Opatz and other collaborators to fill this knowledge-gap. Using a large array of experimental methods and developing a number of new analysis routines, we investigated the effects of common anesthetics on the hippocampus at levels ranging from synapses to cellular ensembles and on memory formation. We were surprised to find very distinct short- and long-term effects between the anesthetics used in this study, despite all of them fulfilling the same hallmarks of general anesthesia. Alterations of brain networks in the hippocampus were also different from those described in the neocortex. In summary, our study suggests that isoflurane should be the anesthetic of choice, if brain processes relevant for memory formation should not be disturbed.” Reference: “Anesthetics fragment hippocampal network activity, alter spine dynamics, and affect memory consolidation” by Wei Yang, Mattia Chini, Jastyn A. Pöpplau, Andrey Formozov, Alexander Dieter, Patrick Piechocinski, Cynthia Rais, Fabio Morellini, Olaf Sporns, Ileana L. Hanganu-Opatz and J. Simon Wiegert, 1 April 2021, PLOS Biology. DOI: 10.1371/journal.pbio.3001146

Recent research on Caenorhabditis elegans suggests that even simple organisms can exhibit basic emotions. This study, which combines behavioral observations and genetic analysis, offers significant insights into the genetic basis of emotions, potentially aiding in the understanding and treatment of human emotional disorders. Insights Into How Short-Term Stimulation Can Alter Sustained Brain Activities and Their Underlying Processes Brain research is one of the most crucial fields in modern life sciences, and “emotion” is one of its major topics. Traditionally, the study of emotions in animals has been a complex area, predominantly examining fear responses in mice and rats. However, since the 2010s, it has been increasingly reported in scientific papers that even crayfish and flies may have brain functions resembling emotions by focusing on several characteristics of their behavior, such as persistence and valence. For instance, when an animal experiences a dangerous situation like being attacked by a predator (a negative valence) even for a short period, the animal’s behavior may be to stay in a safe place, ignoring normally attractive smells of food even if hungry, for a certain length of time (persistence), which can be regulated by a primitive form of emotion. However, the details of these fundamental “emotion mechanisms” remain largely undisclosed. Research on Emotions in Roundworms An international research team from Nagoya City University (Japan) and Mills College at Northeastern University (USA) has revealed the possibility that the roundworm Caenorhabditis elegans possesses basic “emotions.” They used the worms because worms have been used for detailed analysis of basic functions such as perception, memory, and even decision-making at cellular and genetic levels. The team initially discovered that when worms are subjected to alternating current stimulation, worms start moving at an unexpectedly high speed. Illustration of behavioral responses of worms to electric stimulus. Credit: Kristina Galatsis Interestingly, the team also found that this “running” response persisted for 1-2 minutes even after the electrical stimulation for a few seconds was terminated. In animals in general, when a stimulus is stopped, the response to that stimulus usually ceases immediately. (Otherwise, the perception of stimuli such as sounds or visual scenes would linger.) Therefore, the reaction of “continuing to run even after the stimulus stops” is exceptional. Behavioral and Genetic Analysis of Emotional Responses in Worms Furthermore, during and after the electric stimulation, the team found that the worms ignore their food bacteria, which provide crucial environmental information. This suggests that while the presence or absence of their food bacteria is usually crucial, the danger posed by electrical shocks, a survival-threatening stimulus, is even more important. In other words, when worms sense the dangerous stimulus of an electrical shock, their highest survival priority is to escape from that location. To achieve this, the brain’s functioning seems to persistently change, including ignoring the usually significant “food” in order to escape danger. This suggests that the phenomenon of “worms continuing to run due to short-term electrical stimulation” reflects basic “emotions.” Implications for Understanding Human Emotions Furthermore, through genetic analysis, particularly leveraging the advantages of worms, the team revealed that mutants unable to produce neuropeptides, equivalent to our hormones, exhibited a longer duration of continuous running in response to electrical stimulation compared to normal worms. This result indicates that the continuous state in response to danger is regulated to end at the appropriate time. Indeed, if we experience excitement or fear that persists for a very long period, it disrupts our daily lives. Therefore, the findings suggest that our emotions, such as “excitement,” “happiness,” or “sadness,” induced by stimuli, may not be naturally destined to fade away with time, but are controlled by an active mechanism involving genes. This study demonstrates that using worms can offer detailed insights into the genetic mechanisms underlying primitive “emotions”. Many of the genes at work in worms are known to have counterparts in humans and other organisms, so studying worms can offer significant clues about the genes involved in the basis of “emotions.” Specifically, conditions like depression, classified as mood disorders, can be interpreted as states where negative emotions are excessively and persistently maintained due to the inability to effectively process experienced stimuli. If novel genes related to emotions are discovered through worm research, these genes could potentially become targets for new treatments of emotional disorders. Reference: “Electric shock causes a fleeing-like persistent behavioral response in the nematode Caenorhabditis elegans” by Ling Fei Tee, Jared J Young, Keisuke Maruyama, Sota Kimura, Ryoga Suzuki, Yuto Endo and Koutarou D Kimura, 18 August 2023, Genetics. DOI: 10.1093/genetics/iyad148 The study was funded by the Japan Society for the Promotion of Science, Grant-in-Aid for Research in Nagoya City University, the National Institutes of Natural Sciences, the Toyoaki Scholarship Foundation, the Japanese Government (MEXT) Scholarship, and the RIKEN Center for Advanced Intelligence Project (to K.D.K).

Illustration of a human hand as it might appear if the new tissue transparency effect proves effective in humans. Currently, the effect has only been tested with animals in a laboratory setting. Note that dyes may be harmful. Always exercise caution with dyes and do not consume directly, apply to people or animals, or otherwise misuse. Credit: Keyi “Onyx” Li/U.S. National Science Foundation Using common food dye, researchers make skin and muscle safely and reversibly transparent. Scientists at Stanford University have developed a groundbreaking technique using food-safe dye to make animal tissues transparent, enhancing the visibility of internal organs. This innovation has potential applications ranging from medical diagnostics to cancer treatment and has shown promising results in both theoretical and practical tests. Groundbreaking New Imaging Technique Unveiled Researchers have developed a new way to see organs within a body by rendering overlying tissues transparent to visible light. The counterintuitive process—a topical application of food-safe dye—was reversible in tests with animal subjects, and may ultimately apply to a wide range of medical diagnostics, from locating injuries to monitoring digestive disorders to identifying cancers. Stanford University researchers published the research ″Achieving optical transparency in live animals with absorbing molecules″ in the September 6, 2024, issue of Science. ″Looking forward, this technology could make veins more visible for the drawing of blood, make laser-based tattoo removal more straightforward, or assist in the early detection and treatment of cancers,″ said Stanford University assistant professor of materials science and engineering Guosong Hong, a U.S. National Science Foundation CAREER grantee who helped lead this work. ″For example, certain therapies use lasers to eliminate cancerous and precancerous cells, but are limited to areas near the skin’s surface. This technique may be able to improve that light penetration.″ Researchers at Stanford University have developed a way to make skin and other tissues transparent using a simple food dye, a reversible technique with potential for revolutionizing internal medicine. In this clip, thin slices of chicken breast become transparent on exposure to the dye FD & C Yellow 5. Credit: U.S. National Science Foundation An Illuminating Solution To master the new technique, the researchers developed a way to predict how light interacts with dyed biological tissues. Those predictions required a deep understanding of light scattering, as well as the process of refraction, where light changes speed and bends as it travels from one material into another. Scattering is the reason we cannot see through our body: Fats, fluids within cells, proteins, and other materials each have a different refractive index, a property that dictates how significantly an incoming light wave will bend. In most tissues, those materials are closely compacted together, so the varied refractive indices cause light to scatter as it passes through. It is the scattering effect that our eyes interpret as opaque, colored, biological materials. The researchers realized if they wanted to make biological material transparent, they had to find a way to match the different refractive indices so light could travel through unimpeded. Close-up macro image of a gloved hand scooping undissolved yellow #5 dye from a glass jar. Credit: Matthew Christiansen/U.S. National Science Foundation Breakthrough With Tartrazine Building upon fundamental insights from the field of optics, the researchers realized dyes that are the most effective at absorbing light can also be highly effective at directing light uniformly through a wide range of refractive indices. One dye the researchers predicted would be particularly effective was tartrazine, the food dye more commonly known as FD & C Yellow 5. It turns out, they were correct: When dissolved into water and absorbed into tissues, tartrazine molecules are perfectly structured to match refractive indices and prevent light from scattering, resulting in transparency. Close-up macro image of syringe injected a solution of yellow #5 dye into white container filled with water. Credit: Matthew Christiansen/U.S. National Science Foundation From Theory to Practice The researchers first tested their predictions with thin slices of chicken breast. As tartrazine concentrations increased, the refractive index of the fluid within the muscle cells rose until it matched the refractive index of the muscle proteins – the slice became transparent. Then, the researchers gently rubbed a temporary tartrazine solution on mice. First, they applied the solution to the scalp, rendering the skin transparent to reveal blood vessels crisscrossing the brain. Next, they applied the solution to the abdomen, which faded within minutes to show contractions of the intestine and movements caused by heartbeats and breathing. The technique resolved features at the scale of microns, and even enhanced microscope observations. When the dye was rinsed off, the tissues quickly returned to normal opacity. The tartrazine did not appear to have long-term effects, and any excess was excreted in waste within 48 hours. The researchers suspect that injecting the dye should lead to even deeper views within organisms, with implications for both biology and medicine. Illustration of skin tissues as they normally appear, with photons scattering as they interact with surrounding materials. Credit: Keyi “Onyx” Li/U.S. National Science Foundation Illustration of skin tissues rendered transparent following saturation by FD & C Yellow 5, including the paths of photons reflecting off un-dyed tissues. Credit: Keyi “Onyx” Li/U.S. National Science Foundation Illustration of skin tissues rendered transparent following saturation by FD & C Yellow 5. Credit: Keyi “Onyx” Li/U.S. National Science Foundation Old Formulas Yield New Window Into Medicine Supported by a range of federal and private grants, the project began as an investigation into how microwave radiation interacts with biological tissues. In exploring optics textbooks from the 1970s and 1980s, the researchers found two key concepts: mathematical equations called Kramers-Kronig relations and a phenomenon called Lorentz oscillation, where electrons and atoms resonate within molecules as photons pass through. Well studied for more than a century, yet not applied to medicine in this way, the tools proved ideal for predicting how a given dye can raise the refractive index of biological fluids to perfectly match surrounding fats and proteins. Graduate researcher Nick Rommelfanger, working under an NSF Graduate Research Fellowship, was one of the first to realize that the same modifications that make materials transparent to microwaves could be tailored to impact the visible spectrum, with potential applications in medicine. Animation depicting the tissue transparency effect and how it might appear if tested with humans in the future. The latter part of the animation shows how photons interact with tissues at the cellular level, both with and without FD & C Yellow 5 saturation. Credit: Keyi “Onyx” Li/U.S. National Science Foundation Harnessing Old Equipment for New Discoveries Transitioning from theory to experimentation, postdoctoral researcher Zihao Ou—the study’s lead author—ordered a number of strong dyes and began the process of meticulously evaluating each for ideal optical properties. Ultimately, the team grew to 21 students, collaborators, and advisors, involving several analytical systems. One that proved critical was a decades-old ellipsometer nestled among newer equipment at the Stanford Nano Shared Facilities, part of the NSF National Nanotechnology Coordinated Infrastructure (NNCI). The ellipsometer is a tool familiar to semiconductor manufacturing, not biology. However, in a possible first for medicine, the researchers realized it was perfect to predict the optical properties of their target dyes. Time-lapse images of blood vessels in the brain just beneath the skull of a sedated mouse, revealed without any surgery, incisions, or damaging the mouse’s bone or skin. By reversibly dyeing the tissues with FD & C Yellow 5 and using a technique called laser speckle contrast imaging, Stanford University researchers observed the blood flow within this living brain. Credit: Stanford University/Gail Rupert/NSF ″Advanced research facilities constantly aim to strike the right balance by providing access to basic tools and expertise while making space for newer, larger, and more powerful instrumentation,″ said NSF Program Officer Richard Nash, who oversees the NSF NNCI. ″While a basic workhorse such as an ellipsometer would rarely make headlines, it nevertheless can play a crucial role when deployed for atypical uses like the case here. Open access to such instrumentation is foundational for making groundbreaking discoveries, as those instruments can be deployed in new ways to generate fundamental insights about scientific phenomena.″ With methods grounded in fundamental physics, the researchers hope their approach will launch a new field of study matching dyes to biological tissues based on optical properties, potentially leading to a wide range of medical applications. ″As an optics person, I’m amazed at how they got so much from exploiting the Kramers-Konig relationship,″ said NSF Program Officer Adam Wax, who has supported Hong’s work. ″Every optics student learns about them, but this team has used the equations to figure out how a strongly absorbing dye can make skin transparent. Using an NSF EAGER grant, Hong was able to step out in a bold new direction, a great example of how fundamental optics knowledge can be used to create new technologies, including in biomedicine.″ Instrumental NSF Support ″NSF′s support played an instrumental role in the success of this work,″ added Hong. ″The NSF CAREER award was my first major funding, and it arrived at a particularly challenging time, during the darkest moments of the pandemic. My lab faced significant difficulties generating data due to the shutdown, and the award was a vital springboard, enabling me to pursue some of our most exciting and innovative projects – including the research that culminated in this Science paper. The flexibility and encouragement from the NSF awards were crucial in keeping me on track and allowed me the freedom to explore new and uncharted territories in my field.″ Please note: The technique described above has not been tested on humans. Dyes may be harmful. Always exercise caution with dyes and do not consume directly, apply to people or animals, or otherwise misuse. For more on this research: Researchers Make Skin Invisible With Common Food Dye Reference: “Achieving optical transparency in live animals with absorbing molecules” by Zihao Ou, Yi-Shiou Duh, Nicholas J. Rommelfanger, Carl H. C. Keck, Shan Jiang, Kenneth BrinsonJr, Su Zhao, Elizabeth L. Schmidt, Xiang Wu, Fan Yang, Betty Cai, Han Cui, Wei Qi, Shifu Wu, Adarsh Tantry, Richard Roth, Jun Ding, Xiaoke Chen, Julia A. Kaltschmidt, Mark L. Brongersma and Guosong Hong, 6 September 2024, Science. DOI: 10.1126/science.adm6869 This research was supported by NSF grants NNCI 1542152 (NNCI), CAREER 2045120, EAGER 2217582, and GRFP 1656518. In addition to NSF, funders supporting the Stanford research included the U.S. National Institutes of Health, the U.S. Air Force Office of Scientific Research, the U.S. Army Long Term Health Education and Training program, and a range of private foundations and institutions.

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